Biodiversity in Agroecosystems (Advances in Agroecology) - CRC Press (1998)

BIODIVERSITY
IN
AGROECOSYSTEMS
Edited by
Wanda W. Collins and

Calvin O. Qualset
© 1999 by CRC Press LLC.

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Project Editor: Maggie Mogck
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Library of Congress Cataloging-in-Publication Data
Biodiversity in agroecosystems / edited by Wanda Williams Collins, Calvin O. Qualset.

p. cm. — (Advances in agroecology)
Includes bibliographical references and index.
ISBN 1-56670-290-9 (alk. paper)
1. Agricultural ecology. 2. Biological diversity. 3. Agricultural systems.
I. Collins, Wanda Williams. II. Qualset, Calvin O. III. Series.
S589.7.B575 1998
630—dc21 98-13056
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Preface
Diversity in biological resources (biodiversity) evolved to fill the innumerable
environmental niches of the Earth. It is manifest in all except the most extreme
global environments. Humans have exploited this diversity and shaped it over mil-
lennia to meet basic needs for survival. Thus, biodiversity has been an integral and
essential feature in the evolution of agriculture. Agrobiodiversity, that part of the
full spectrum of diversity on which humans directly depend for food and fiber,
represents plants, animals, and microbes that are greatly modified by the domesti-
cation process to the extent that many of them cannot survive in the environments
of their progenitors.
Ancient farmers knew well the necessity of having different crops or animal
breeds and different varieties of each crop or breed in case of failure of one or the
other for various reasons. They also explored and utilized the rich diversity of flavors,
textures, and aromas available in their plant and animal species.
The management of agrobiodiversity is temporally and spatially integrated into
agroecosystems. How agroecosystem management affects ecosystems in general is
a subject of intensive research, debate, and controversy. Both intensive and extensive
agricultural systems are essential to the sustainability of the global human popula-
tion. In turn, these systems are dependent upon the existence of a rich reservoir of
agrobiodiversity and its use in ways that complement and enhance biodiversity in
its natural state. As is evident throughout this book, there are examples of severe
negative impacts of agricultural activities on biodiversity, mainly through degrada-
tion of natural habitats. Biodiversity losses are becoming recognized and, through
research and practice, are slowly being mitigated. At the same time, there are
examples of agriculture practices that enhance biodiversity and one of the great
challenges for the next century will be to discover additional ways for achieving
complementarity of food, fiber, and energy production and biodiversity conservation.
Today’s farmers, using both traditional and high-technology methods, also under-
stand that the continued success of their agriculture depends upon conserving,
maintaining, and using the diversity that is so under threat now. It is also important
that conservationists and policy makers have a thorough understanding of agricul-
tural processes so they can discover practical ways to facilitate both agricultural
production and natural resource conservation.
This book is designed to emphasize just how important biodiversity is to agri-
culture. The various chapters point out the positive effects of agriculture on helping
to conserve and protect the diversity on which it depends. The effects of agricultural
growth, genetic uniformity, and dependence on high levels of external inputs are
also highlighted. The book is intended to present the reader with a broad view of
the interplay of biodiversity and agriculture with chapters addressing soil microbes,
insects, plants, animals, rangelands, and agro-forestry. Other chapters discuss the
efforts to conserve, maintain, and effectively use plant and animal diversity most
important to agriculture. Still other chapters delve into the value of genetic diversity
to agriculture and the interaction with surrounding habitats and species. Finally, the
integration of new biotechnologies into traditional and industrial agricultural systems

can have great impact on the quality of biodiversity and help shape strategies for its
conservation.
We believe that sharing this knowledge and these experiences by some of the
world’s most knowledgeable experts will provide readers with a broad appreciation
and heightened awareness of the stakes involved in the future preservation of these
natural resources.
Wanda W. Collins
Calvin O. Qualset

The Editors
Wanda W. Collins, Ph.D., currently is the Deputy Director General for Research
at the International Potato Center in Lima, Peru. She is the author of more than 40
journal articles on genetics and plant breeding, several book chapters, and numerous
extension and popular publications.
Dr. Collins is a graduate of North Carolina State University where she received
her Ph.D. in both Plant Pathology and Genetics in 1976. She joined the school’s
Department of Horticultural Science that same year and became a full professor in
1986. She has been doing research on vegetable breeding and genetics (sweetpotato
and potato) since 1976 and traveled to Peru to research the genetic resources of the
Ipomoea species (wild relatives of the sweetpotato). Between 1996 and 1998, Dr.
Collins also spent over 2 years on study leave at the World Bank in Washington,
D.C. as Agricultural Research Advisor, Environmentally Sustainable Development
Vice Presidency.
She is one of the originators and the first chair of the Sweetpotato Crop Genetics
Committee which advises the U.S. Germplasm System on sweetpotato germplasm.
Her many memberships and contributions to the field of horticulture include Chair
of the Board of Trustees for the International Plant Genetic Resources Institute,
member of the Executive Board for the American Society of Horticultural Science,
and as the current President of the American Society of Horticultural Science.
Calvin O. Qualset, Ph.D., is Director of the Genetic Resources Conservation

Program of the Division of Agriculture and Natural Resources at the University of
California, Davis. Before retiring as a professor in 1994, he held the positions of
Chair of the Department of Agronomy and Range Science and Assistant Dean for
Plant Sciences and Pest Management at UC Davis. He also served as Assistant
Professor at the University of Tennessee, Knoxville from 1964 to 1967 before
moving to California.
He is active in developing biological conservation strategies and conducts
research on plant genetic resource conservation and genetics. He serves on the Board
of Trustees of the American Type Culture Collection as a representative of the
Genetics Society of America. He coordinates the International Triticeae Mapping
Initiative and is Principal Investigator on McKnight Foundation and BARD-funded
projects in Mexico and Israel on in situ conservation of crop plants and genetic
mapping of wheat. He has served on numerous reviews of international centers for
the Consultative Group for International Agricultural Research, of university depart-
ments, and of current issues in agriculture and biology for the National Science
Foundation and National Research Council.
Dr. Qualset held Fulbright Fellowships to Australia and Yugoslavia and served
as president of the American Society of Agronomy and of the Crop Science Society
of America. He has received citations for work in plant breeding from the U.S.
National Council of Commercial Plant Breeders and the Mexican National Institute
of Agriculture and Forestry. He has authored more than 190 scientific papers and
developed numerous varieties of wheat, oat, and triticale.

Contributors
Miguel A. Altieri Joel I. Cohen
Center for Biological Control Project Manager, Intermediary
ESPM-Division of Insect Biology Biotechnology Service
201 Wellman – 3012 International Service for National
University of California at Berkeley Agricultural Research (ISNAR)
Berkeley, CA 94720-3112 The Hague, The Netherlands
([email protected]) ([email protected])
Stuart S. Bamforth Wanda W. Collins (Editor)

Department of Ecology, Evolution, and Deputy Director General for Research
Organismal Biology International Potato Center
Tulane University Apartado 1558
New Orleans, LA 70118-5698 Lima 12, Peru
([email protected])
Mary E. Barbercheck
Department of Entomology Cornelis de Haan
North Carolina State University World Bank
Raleigh, NC 27695-7634 1818 H Street N.W.
([email protected]) Washington, D.C. 20433
([email protected])
Gary W. Barrett
Institute of Ecology Michael I. Goldstein
University of Georgia Department of Wildlife and Fisheries
Athens, GA 30602-2202 Science
([email protected]) Texas A&M State University
210 Nagle Hall
Terry A. Barrett College Station, TX 77843-2258
Institute of Ecology ([email protected])
University of Georgia
Athens, GA 30606-2202 Douglas Gollin
([email protected]) Department of Economics
Fernald House, Williams College
Harvey W. Blackburn Williamstown, MA 01267 and
Research Leader Affiliate Scientist, Economics Program
USDA Sheep Experiment Station Centro Internacional de Mejoramiento
H.C. 62, Box 2010 de Maiz y Trigo (CIMMYT)
Dubois, ID 83423 Lisboa 27, Apdo. Postal 6-641,
([email protected]) CP 06600 Mexico, D.F.
([email protected])
Lara M. Coburn
Department of Wildlife and Fisheries Keith Hammond
Science Animal Production and Health Division
Texas A&M State University Food and Agriculture Organization of
210 Nagle Hall the United Nations
College Station, TX 77843-2258 00100 Rome, Italy

Geoffrey C. Hawtin Clara I. Nicholls
Director General Department of Entomology
International Plant Genetic Resources University of California at Davis
Institute (IPGRI) Davis, CA 95616
Via delle Sette Chiese 142 ([email protected])
Rome, Italy
([email protected]) John David Peles
Savannah River Ecology
Ann C. Kennedy Laboratory
USDA-ARS Drawer E
Pullman, WA 99164-6421 Aiken, SC 29806
Thomas E. Lacher, Jr. Keith S. Pike

Department of Wildlife and Fisheries Irrigated Agriculture Research and
Science Extension Center
Texas A&M State University Washington State University
210 Nagle Hall Prosser, WA 99350
College Station, TX 77843-2258 ([email protected])
([email protected])
Jon K. Piper
Roger R. B. Leakey Department of Biology
Institute of Terrestrial Ecology, Bush Bethel College
Estate North Newton, KS 67117
Penicuik, Midlothian EH26 0QB ([email protected])
Scotland, U.K.
([email protected]) Calvin O. Qualset (Editor)
Director, Genetic Resources
Helen W. Leitch Conservation Program
International Livestock Research University of California at Davis
Institute Davis, CA 95616
Box 30709 ([email protected])
Nairobi, Kenya
([email protected]) Robert E. Rhoades
Department of Anthropology
Virginia D. Nazarea University of Georgia
Department of Anthropology Athens, GA 30602
University of Georgia ([email protected])
Athens, GA 30602
([email protected]) R. Douglas Slack
Department of Wildlife and Fisheries
Deborah A. Neher Science
Department of Biology Texas A&M State University
University of Toledo 210 Nagle Hall
Toledo, OH 43606 College Station, TX 77843

Melinda Smale Neil E. West
Economics Program Department of Rangeland Resources
Centro Internacional de Mejoramiento Utah State University
de Maiz y Trigo (CIMMYT) Logan, UT 84322
Lisboa 27, Apdo. Postal 6-641 ([email protected])
CP 06600 Mexico, D.F
([email protected])
Petr Starý
Institute of Entomology
Czech Academy of Sciences
Branisovska 31,
370 05 Ceské Budejovice
Czech Republic

Contents
Chapter 1
Microbial Diversity in Agroecosystem Quality
Ann C. Kennedy
Chapter 2
Soil Microfauna: Diversity and Applications of Protozoans in Soil
Stuart S. Bamforth
Chapter 3
Diversity and Function of Soil Mesofauna
Deborah A. Neher and Mary E. Barbercheck
Chapter 4
Uses of Beneficial Insect Diversity in Agroecosystem Management
Petr Starý and Keith S. Pike
Chapter 5
Biodiversity, Ecosystem Function, and Insect Pest Management in
Agricultural Systems
Miguel A. Altieri and Clara I. Nicholls
Chapter 6
Livestock and Biodiversity
Harvey W. Blackburn and Cornelis de Haan
Chapter 7
Managing for Biodiversity of Rangelands
Neil E.West
Chapter 8
Agroforestry for Biodiversity in Farming Systems
Roger R. B. Leakey
Chapter 9
The Role of Agroecosystems in Wildlife Biodiversity
Thomas E. Lacher, Jr., R. Douglas Slack, Lara M. Coburn, and
Michael I. Goldstein
Chapter 10
Natural Systems Agriculture
Jon K. Piper

Chapter 11
Managing Agroecosystems as Agrolandscapes: Reconnecting
Agricultural and Urban Landscapes
Gary W. Barrett, Terry A. Barrett, and John David Peles
Chapter 12
Local Management of Biodiversity in Traditional Agroecosystems
Robert E. Rhoades and Virginia D. Nazarea
Chapter 13
Valuing Genetic Diversity: Crop Plants and Agroecosystems
Douglas Gollin and Melinda Smale
Chapter 14
Conserving and Using Crop Plant Biodiversity in Agroecosystems
Wanda W. Collins and Geoffrey C. Hawtin
Chapter 15
Implementing the Global Strategy for the Management of Farm
Animal Genetic Resources
Keith Hammond and Helen W. Leitch
Chapter 16
Agroecosystem Quality: Policy and Management Challenges for
New Technologies and Diversity
Joel I. Cohen

CHAPTER 1
Microbial Diversity in
Agroecosystem Quality
Ann C. Kennedy
CONTENTS
Introduction
Taxonomic Diversity
Functional Diversity
Soil Microorganisms in Agroecosystems
Rhizosphere
Microbial Impacts on Agroecosystems
Plant Growth
Nutrient Cycling
Soil Structure
Agroecosystem Impacts on Microorganisms
Cropping Systems
Tillage
Potential Application of Microbial Indicators
Conclusion
Acknowledgments
References
INTRODUCTION
Agroecosystems comprise 30% of Earth’s surface (Altieri, 1991). Microbial

diversity is one factor that controls agroecosystem productivity and quality. Biolog-
ical diversity studies generally have been centered around plants, animals, and insect

Table 1 Justification for Investigations of Microbes and Microbial Diversity
Microbes are an important source of knowledge about life strategies and limitations.
Microbes are key to understanding evolutionary history.
The untapped diversity of microorganisms is a resource for new genes and organisms of value
to biotechnology.
Microbes and microbial diversity patterns can be used as indicators of environmental changes.
Microbial communities are key to understanding biological interactions.
Microbes play a key role in conservation and restoration biology of higher organisms.
Microbes are critical to sustainability.
Source: Microbial Diversity Research Priorities, American Society of Microbiology, Washington,
D.C., 1994. With permission.
species with little attention being given to microorganisms. Without microorganisms

and their biochemical processes, life on Earth would not be possible (Price, 1988).
Microbial diversity can directly influence plant productivity and diversity by influ-
encing plant growth and development, plant competition, and nutrient and water
uptake. Microbes provide sources of genetic material; in addition, they can be used
as indicators of environmental change (American Society of Microbiology, 1994;
Table 1).
Microorganisms are key to understanding biological interactions. They play a
role in conservation and restoration of agroecosystems and are essential to quality
considerations. Soil organisms constitute a large, dynamic nutrient source in all
ecosystems and play a major role in residue decomposition and nutrient cycling
(Smith and Paul, 1990). In addition, microorganisms are responsible for chemical
changes in the soil, such as the accumulation of soil organic matter, dinitrogen
fixation, and other changes in soil properties that may affect plant growth. When a
disturbed system begins to recover, it is not only due to plant recolonization, but is
also dependent upon the action of microbes in promoting favorable soil conditions
and fertility.
We are unaware of the true extent or dimension of the diversity of soil microbes,
although molecular investigations suggest that populations in soil are greater than
we are able to understand with cultural techniques (Holben and Tiedje, 1988; Torsvik
et al., 1990). Microbial diversity is critical to ecosystem functioning as a result of
the myriad of processes for which microbes are responsible, such as decomposition
and nutrient cycling, soil aggregation, and pathogenicity. It is necessary for us to
increase our knowledge of biotic and functional diversity to understand better the
desired makeup of microbial communities and the optimum management practices
for an agroecosystem.
Anthropogenic influences may affect ecosystem functioning and diversity. Some
of the most dramatic examples of ecosystem disturbances are occurring as a result
of soil erosion from agroecosystem perturbations. These disturbances may be leading
to progressive declines in biological, chemical, and physical stability in ecosystems.
Continued disturbance of these areas may cause massive changes in global carbon
cycling, resulting from significant organic carbon loss from the terrestrial environ-
ment. Microorganisms are highly sensitive to disturbance, such as those introduced
by agriculture (Elliott and Lynch, 1994), and they may function as early warning
indicators of changes in quality. Fluxes in microbial diversity and functional diversity

may contribute greatly to the understanding of soil quality and the development of
sustainable agroecosystems (di Castri and Younes, 1990; Hawksworth, 1991a; Tho-
mas and Kevan, 1993). Soil organisms are useful in classifying disturbed or con-
taminated systems, since diversity can be affected by minute changes in the ecosys-
tem. The use of microorganisms and their functioning for examination of
environmental stress and declining biological diversity needs to be exploited for the
benefit of agroecosystems (Office of Technology Assessment, 1987). The purpose
of this chapter is to explore the issue of microbial diversity in agroecosystems and
its influence on the quality of that agroecosystem.
Taxonomic Diversity
The number of known microbial species is estimated to be over 110,000, but

only a fraction of these species is identified and even fewer are being studied or are
in culture collections (Hawksworth, 1991b). By one estimate, only 3 to 10% of
Earth’s microbial species have been identified, leaving a vast portion of that biota
unknown and therefore unstudied (Hawksworth, 1991a). According to other esti-
mates, the number of microbial species may be as many as 1 million organisms
(American Society for Microbiology, 1994). Even with conservative estimates, the
microbial populations and their diversity require more investigation. Microbial com-
munities are highly diverse and thought to exhibit even greater diversity than that
seen in higher orders of organisms (Torsvik et al., 1990; Ward et al., 1992). Obvi-
ously, we are aware of only a miniscule portion of the total potential of the system.
Within the soil microbial population, there is a wealth of genetic information waiting
to be discovered.
Functional Diversity
Functional diversity and taxonomic diversity are often two vastly different mea-
surements. Functional diversity includes the magnitude and capacity of soil inhab-
itants that are involved in key roles. These processes are selected to represent
biologically meaningful processes, such as carbon or nitrogen cycling, decomposi-
tion of various compounds, and other transformations (Zak et al., 1994). Taxonomic
diversity is determined by culturability and isolation of species, and may represent
20% or less of the microbes present and active in soil. Speciation relies on charac-
terization of known phenotypic or genetic characterizations that may not be present
in soil and possibly have no bearing on soil processes (Lee and Pankhurst, 1992).
Microbial diversity indexes have been used to describe the status of microbial
communities and their response to natural or human disturbances. Microbial diversity
indexes can function as bioindicators of the stability of a community and can be
used to describe the ecological dynamics of a community and the impact of stress
on that community (Mills and Wassel, 1980; Atlas, 1984). Chemically stressed or
heavy metal–stressed soils were found to decrease in microbial diversity depending
on the type of chemical applied (Atlas, 1984; Reber, 1992). A factor limiting the
greater use of these indexes is the absence of detailed information on the microbial
species composition of soil environments (Torsvik et al., 1990).

The ability of an ecosystem to buffer the effects of extreme disturbances may
depend in part on the diversity of the system (Elliott and Lynch, 1994). It may be
important to monitor diversity as an indicator of change or in response to a stress.
The extinction rate of species within a system may be an important indicator of the
status of the system and critical in determining the level of diversity necessary to
maintain an agroecosystem. The actual numbers of species and species composition
may not be as important as the flux of species within a system and the functioning
of those individuals within that system.
Diversity indexes can be used to indicate the effect of disturbance; however,
greater diversity may not always be desirable. Greater diversity should not be equated
with a more stable system; rather, the changes in diversity with management may
be more informative of the status of a soil microbial community. Diversity issues
in understanding quality of soil may not be enlightening unless the functions of the
system are taken into account. Basic shifts in large groups in an ecosystem may
indicate a change, but may not be able to address functioning of that altered system.
Microbial communities and their processes need to be examined, not only in relation
to the individuals that make up the community, but also with regard to the effect of
perturbations or environmental stresses on those communities.
SOIL MICROORGANISMS IN AGROECOSYSTEMS
The soil is full of microorganisms such as bacteria, actinomycetes, fungi, algae,

viruses, and protozoa. Microbes are classified in categories based on their utilization
of carbon and energy and their nutrient requirements (i.e., phototrophs, chemotrophs,
autotrophs, heterotrophs, or lithotrophs). This provides limited information, since
many situations overlap and refinement is needed (Alexander, 1977; Walker, 1992;
Zak et al., 1994). Bacteria and fungi, as primary decomposers in the cycling of
nutrients, occupy a critical position in the soil food web. Of all nutrient cycling, 90
to 95% passes through these two groups of organisms to higher trophic levels. Thus,
the function and possibly the diversity of bacteria and fungi will be a large factor
in determining the quality of agroecosystems (Lynch, 1983).
Bacteria and actinomycetes are the most numerous of microbial groups in soils,
but because of their small size, 1 to 10 µm, they only account for 50% of the total
biomass in soil (Alexander, 1977). Bacteria are found in soil at populations of 104
to 109 cells g–1 soil. As a group they are diverse metabolically and use many different
sources of energy and carbon. Bacteria play an important role in the breakdown of
organic material and nutrient cycling. Most natural and xenobiotic compounds can
be broken down by soil microflora with few compounds becoming recalcitrant (Dorn
et al., 1974). Some bacteria have the potential for nitrogen fixation (Sprent, 1979)
or methane production or utilization (Jones, 1991). Denitrification and sulfate reduc-
tion involve a variety of facultative and obligate anaerobic bacteria (Tiedje et al.,
1984). Nitrification and sulfur oxidation, on the other hand, are the result of the
activity of a limited number of genera of aerobic autotrophic bacteria (Belser and
Schmidt, 1978; Bock et al., 1989).

Fungi are less diverse than bacteria and less numerous in soil at 104 to 106
propagules g–1 soil. Fungi however, are responsible for up to 70% of the biomass
(Lynch, 1983). Fungi are found in soil, in associations with plant roots, or as
saprophytes on detrital material (Swift and Boddy, 1984). Fungi can withstand
adverse soil conditions better than other microorganisms, and they survive at lower
water potentials than bacteria (Papendick and Campbell, 1975). Hyphal strands allow
fungi to overcome the environmental constraints of low moisture and depleted
nutrients by translocation of water and nutrients. Also, several fungi excrete organic
acids that solubilize otherwise unavailable nutrients (Sollins et al., 1981). Fungi are
active decomposers of cellulose, lignin, and other organic materials. The products
of decomposition are then released and used by other organisms, especially bacteria.
Many fungi are plant pathogens, yet some form beneficial relationships with plant
roots (e.g., mycorrhizae).
Algae are most numerous in surface soil and found at populations of 102 to 106
cells g–1 soil. In some agricultural systems, algae contribute to nitrogen cycling by
nitrogen fixation or to soil stabilization (Metting and Rayburn, 1983). Protozoa
inhabit soil at populations between 103 and 105 cells g–1 soil. These organisms are
major predators of bacteria and thus regulate bacterial populations (Opperman et
al., 1989).
Rhizosphere
Plants may be an important selective force for the diversity of rhizosphere

populations of bacteria and fungi through their influence on soil nutrients. Rhizo-
sphere soil can be defined as that volume of soil adjacent to and influenced by plant
roots (Metting, 1993). It is a region of intense microbial activity because of its
proximity to plant root exudates, making rhizosphere microbial communities distinct
from those of bulk soil (Curl and Truelove, 1986; Whipps and Lynch, 1986). Micro-
bial activity is stimulated in this area because of the nutrients provided by the root
or germinating seed (Rouatt and Katznelson, 1961). Microbial populations and
functioning in an agroecosystem are influenced by the root and the soil environment,
including mineral and organic material. The aboveground plant community can
influence microbial spatial heterogeneity in soil. Those microorganisms that respond
to root exudates or related substrates will dominate the rhizosphere. Decaying root
systems also function as a source of nutrients for the surrounding microorganisms
(Swinnen et al., 1995). Microbial populations decrease with distance from the roots
(Yeates and Darrah, 1991).
Bacteria account for the largest number of inhabitants of the rhizosphere. Gram-
negative, non-spore-forming rods with simple nutritional requirements are stimulated
more by roots than are coccoid forms and Gram-positive, spore-forming rods (Curl
and Truelove, 1986). The composition of the plant community may influence the
diversity of the microbial community due to the variability in chemical composition
of the exudates (Christensen, 1989).
The development of rhizosphere microbial communities also is influenced by dif-
ferent plant species (Rovira, 1956), plant phenology (Smith, 1969), and environmental

factors influencing plant growth (Rovira, 1959; Vancura, 1967; Martin and Kemp,
1980). Bacterial and fungal abundance in the rhizosphere is influenced by the nutrient
status of both plant and soil. The percent mycorrhizal cover on roots of Plantago
lanceolata was positively correlated with leaf nitrogen and phosphorus, while the
percent cover of bacteria and other fungi was negatively correlated with phosphorus
(Newman et al., 1981). Nitrogen fertilization increased numbers of fungi and Gram-
negative bacteria in a rice rhizosphere (Emmimath and Rangaswami, 1971). It may
be difficult to separate the effects of soil nutrients on rhizosphere populations from
effects involved with increased or altered root exudation of organic compounds.
Grasses grown in monoculture can modify nitrogen availability (Wedin and Tilman,
1990), and it has been hypothesized that such changes in soil nitrogen availability
influenced by plant species affects the composition of vesicular arbuscular mycor-
rhizae fungal communities (Johnson et al., 1992b).
MICROBIAL IMPACTS ON AGROECOSYSTEMS
Microbes impact agroecosystems through a large list of functions for which they
are responsible (Table 2). Soil humus formation, cycling of nutrients, and building
soil tilth and structure (Lynch, 1983; Wood, 1991) are distributed among a large
number of different genera and species. Microorganisms are responsible for many
transformations in soil related to plant nutrition and health. The majority of soil
microbes are beneficial to plant growth, but they need to be managed effectively
(Lynch, 1983). Potential harmful effects from soil microorganisms include plant
disease, production of plant-suppressive compounds, and loss of plant-available
nutrients. Specific microorganisms can be manipulated to produce beneficial effects
for agriculture (Lynch, 1983), for example, rhizobia to increase plant available
nitrogen (Sprent, 1979), mycorrhizal associations to enhance nutrient uptake
(Mohammad et al., 1995), or biological control of plant pests to reduce chemical
inputs (Cook and Baker, 1983; Kennedy et al., 1991).
Beneficial soil bacteria can enhance plant performance by an increase in mineral
solubilization (Okon, 1982), dinitrogen fixation (Albrecht et al., 1981), the produc-
tion of hormones (Brown, 1972), and the suppression of harmful pathogens (Chang
and Kommendahl, 1968; Cook and Baker, 1983). The symbiotic relationship between
bacteria and legumes is one of the most widely studied and applied plant–microbial
Table 2 Several Functions of Soil Microbes

Decomposition of organic residues with release of nutrients
Formation of beneficial soil humus by decomposing organic residues and through synthesis of
new compounds
Release of plant nutrients from insoluble inorganic forms
Improved plant nutrition through mycorrhizal relationships which are symbiotic relationships
between fungi and plant roots
Transformation of atmospheric dinitrogen to plant-available N
Improvement of soil aggregation, aeration, and water infiltration
Antagonistic action against insects, plant pathogens, and weeds (biological control)

interactions (Sprent, 1979). The bacterium Rhizobium forms nodules on the roots
of the legume plant, takes dinitrogen from the air, and transforms it to plant-available
nitrogen (i.e., NH4+, NO3–). The plant provides nodules and photosynthate for the
bacteria, while the bacteria give the plant the nitrogen it needs. Inoculation of
legumes with dinitrogen-fixing Rhizobium can add appreciable amounts of nitrogen
to the soil. The distribution and diversity of specific strains of dinitrogen-fixing
bacteria vary with environmental conditions (Turco and Bezdicek, 1987; Hirsch et
al., 1993). The plant community can influence the presence or absence of specific
strains of Rhizobium, and thus impact the diversity of this group of microorganisms
(Strain et al., 1994). The interaction involving mycorrhizal fungi and rhizobia may
further affect the host plant by increasing nitrogen and phosphorus nutrition (Allen,
1992). These interactions are specific (Molina et al., 1992), further illustrating the
complexity of the plant–microbe interaction and the changes in diversity of various
microbial groups that can affect plant growth or impact other soil features.
Plant-suppressive bacteria reduce seed germination and delay plant development
by the production of phytotoxic substances (Woltz, 1978; Suslow and Schroth, 1982;
Alstrom, 1987; Schippers et al., 1987). Pathogenic fungi greatly reduce the survival,
growth, and reproduction of plants (Shipton, 1977; Bruehl, 1987; Burdon, 1987), while
beneficial mycorrhizal fungi can enhance plant growth by increasing nutrient (Fitter,
1977; Hall, 1978; Rovira, 1978; Ocampo, 1986) and water uptake (Tinker, 1976).
Mycorrhizal fungi have been found in a vast majority of plant communities
(Allen and Allen, 1990) growing in association with 90% of terrestrial plants exam-
ined (Harley and Smith, 1983). Mycorrhizae are nonpathogenic fungi that form
symbiotic associations with plant roots. The diversity of this group of organisms is
vast, yet is not well studied. These associations are highly specific and indicate the
diversity of this group of microorganisms (Molina et al., 1992). Mycorrhizal asso-
ciations have been shown to be of greatest importance in stressed environments,
phosphorus-deficient soils, eroded sites, and acidic or reclaimed lands (Harley and
Smith, 1983; Barea, 1991). This association may be key in plant productivity and
nutrient cycling (Barea, 1991; Allen, 1992). Mycorrhizal associations are enhanced
by crop rotation and management practices favoring minimum disturbance.
Plant Growth
Interactions between plants and rhizosphere microbes may play a critical role in
the outcome of plant competition. Competitive interaction among plants may also
be important for the development of rhizosphere soil communities. Microbes affect
nutrient uptake (Tinker, 1976; Okon, 1982). Microorganisms may play important
roles affecting plant competition acting as mutualistic or pathogenic plant associates
(Allen and Allen, 1990). Rhizosphere microorganisms may affect plant growth
directly (Woltz, 1978; Suslow and Schroth, 1982; Gaskins et al., 1985; Alstrom,
1987; Schippers et al., 1987) or indirectly by their effects on each other and the
microscale alteration of soil nutrients. Plant performance may also be affected by
competitive interactions between adjacent plants (Goldberg and Fleetwood, 1987;
Gurevitch et al., 1990; Goldberg and Landa, 1991).

Another example of microbial diversity influencing plant growth is the investi-
gation of deleterious rhizobacteria, which were discovered in the early 1980s. Inves-
tigations of deleterious rhizobacteria have led to changes in management practices
for many crops (Schippers et al., 1987) and may lead to biological control of weed
species (Kremer et al., 1990; Kennedy et al., 1991). Deleterious rhizobacteria that
specifically inhibit various grass weeds, but do not affect the crop, have been isolated
from soil (Kennedy et al., 1991; 1992; Kennedy, 1994) and inhibit plant growth by
the production of plant-suppressive compounds (Tranel et al., 1993). These bacteria
are excellent biological control agents, in part, because they are aggressive colonizers
of the roots and residue, often constituting up to 95% of the total pseudomonads on
the plant root (Stroo et al., 1988; Kennedy et al., 1992). Biocontrol is critical to
sustainable agriculture systems, but a greater understanding of soil microbes and
their ecology is needed before biological control can be implemented. There is a
wealth of genetic material contained within the soil that may have potential in
biotechnology programs; thus, diversity investigations will benefit more than just
one area of science (Malik and Claus, 1987; Bull et al., 1992).
Nutrient Cycling
Soil microorganisms constitute a large dynamic source and sink of nutrients in

all ecosystems and play a major role in plant litter decomposition and nutrient cycling
(Smith and Paul, 1990; Collins et al., 1992; Cambardella and Elliott, 1992). Microbes
break down complex compounds in organic residue to simpler, smaller compounds,
as well as recalcitrant compounds. This recycling into compounds of various com-
plexity provides substrate for other microbes, further sustaining the diversity of soil
microbes. Organic matter, in various forms of decay, improves soil physical prop-
erties, increases water-holding capacity, increases nutrient availability, and acts as
a cementing agent for holding soil particles together. Organic matter can be main-
tained by incorporation of crop residues, crop rotation, and addition of animal and
green manures when possible. Addition of organic matter aids in ensuring a produc-
tive soil and stimulates plant growth by providing food for microorganisms.
Soil Structure
Microbes play a major role in the formation of soil structure (Lynch and Bragg,
1985; Tisdall, 1991). Fungi and actinomycetes produce hyphal threads that bind soil
particles together. Extracellular polysaccharides produced by bacteria and fungi bind
soil particles together, assisting in building soil structure. Humic materials from
microbial action form organic matter/clay complexes. This action reduces erosion,
allows for good water infiltration, and maintains adequate aeration of the soil. Soil
aggregation can be increased by the addition of residues resulting in additional
microbial activity (Gilmour et al., 1948). The carbon and nitrogen pools delimit
microbial biomass and decomposition rates and polysaccharide production (Knapp
et al., 1983). The ability of fungi and bacteria to influence aggregation varies with
species and is substrate dependent (Aspiras et al., 1971). Limited nitrogen in soil
solution reduces biomass while increasing polysaccharide production, which can

lead to increased aggregation (Elliott and Lynch, 1984). Soil stability may be man-
aged by addition of different amendments to stimulate microbial activity (Jordahl
and Karlen, 1993).
AGROECOSYSTEM IMPACTS ON MICROORGANISMS
Microbial diversity considerations need to be included in soil quality investiga-

tions of agricultural lands (Kennedy and Papendick, 1995). Soil quality is critical
to the functioning of any ecosystem (Papendick and Parr, 1992). The quality of a
soil can greatly impact land use, sustainability, and productivity. Soil quality, the
inherent characteristic of a soil, comprises physical, chemical, and biological prop-
erties. These properties may be altered through intensive management practices.
Reduction in aboveground plant diversity that occurs with severe disturbance, such
as tillage, overgrazing, and pollutants, may decrease microbial diversity
(Gochenauer, 1981; Boddy et al., 1988; Christensen, 1989).
Cropping Systems
Microbial communities in agroecosystems change with management history. In

continuous cropping systems, cycling of pathogens and antagonists of pathogens
and changes in crop yields can often be seen due to alteration in the disease pressure
over time. An example of this is the decline of a disease of wheat called take-all
decline that is the result of a change in the soil microbial community. The microbial
community shifts to favor growth of antagonists of the pathogen Gaeumannomyces
graminis var. tritici which results in decline of the disease (Cook, 1981). An example
of the use of soil microbial diversity is the emerging area of biological control of
plant pathogens (Cook and Baker, 1983). Microbes have the potential to be used in
biological control, which is the suppression of one pest by using its natural predator
or antagonist. Biocontrol can be used to control insects, pathogens, and weeds by
either lowering the populations of the pest or by reducing the impact of the pest
(DeBach, 1964). Microbes function as a direct delivery system for the natural
pesticide they produce. Bacteria and fungi that produce different types of antibiotics
can be used to control many plant pathogens (Cook and Baker, 1983).
Crop rotation is a key component in sustainable systems because it enhances
beneficial microbes, interrupts the cycle of pathogens, and reduces weed populations.
Legumes in rotation supply symbiotically fixed nitrogen to the system, aid in main-
taining proper water status, and reduce pathogen load. Studies have shown the
positive effects of crop rotation on crop growth, attributing this to changes in the
microbial community composition (Shipton, 1977; Cook, 1981; Johnson et al.,
1992a). Continuous monocropping led to changes in the soil community which
increased the pathogen load and reduced barley growth when compared with grains
in multiple-crop rotation (Olsson and Gerhardson, 1992). The populations and
aggressiveness of pathogens can be altered with crop rotation, illustrating the changes
in microbial diversity and function due to management (El Nashaar and Stack, 1989).
In a long-term study, Cochliobolus sativus, a pathogen of spring wheat, was found

in higher numbers and individual isolates exhibited greater aggressiveness or ability
to cause severe disease in continuous wheat rotation, when compared with wheat in
a 3-year rotation.
Tillage
In a study of the diversity of prairie and cultivated soils, diversity indexes were
greater in disturbed or cultivated systems when compared with grassland (Kennedy
and Smith, 1995). The increase in diversity with disturbance indicates a change in
the microbial community to one that exhibited a greater range of substrate utilization
and stress resistance. Soil microorganisms may affect plant growth and may influence
plant competition. In turn, plants may act as a selective force for rhizosphere
microbial populations through their influences on soil nutrients.
The ecology of root–microbe interactions after minimum tillage practices is
vastly different from that after extensive plowing to prepare the seedbed. The
changes in the physical and chemical properties of the soil resulting from tillage
greatly alter the matrix-supporting growth of the microbial population. Within a
given soil, there is considerable variation in the composition of the microbial
community and diversity with depth in the profile. In no-till agricultural systems,
microbial activities differed drastically with depth, with the greatest microbial
activity occurring near the no-till surface; in the tilled system, activities were more
evenly distributed throughout the plow layer (Doran, 1980). The composition of the
microbial community influenced the rate of residue decomposition and nutrient
cycling in both no-till and conventionally tilled systems (Beare et al., 1993). Decom-
position in the no-till system was dominated by fungi, while the bacterial component
was found to be responsible for a greater portion of the decomposition of residue
in conventionally tilled systems. These studies illustrate the alteration of the makeup
of the microbial communities and possibly the diversity of basic microbial groups
with changes in management systems.
POTENTIAL APPLICATION OF MICROBIAL INDICATORS
The diversity of the microbial community, as well as the functions within the
community, affect the stability and resilience of the soil system. Neither a higher
nor lower degree of diversity in a system can be said to be better or worse; however,
changes in the activity or community structure may influence the quality of the soil.
The microbial biomass has recently been used as a sensitive indicator of manage-
ment-induced changes (Doran, 1987; Powlson et al., 1987; Visser and Parkinson,
1992). Researchers have found discrepancies in microbial biomass responses to
perturbations such as tillage. Kennedy and Smith (1995) found increased diversity
with tillage. Utilizing microbial characteristics may better forecast change in soils
since they respond to perturbations more rapidly than other indicators (Kennedy and
Papendick, 1995).
Soil quality measures have recently become important in system comparisons
of management options. Integral to soil quality assessment are measures of the

microbial community. Microbiological properties can identify changes in the overall
soil quality before changes occur in certain physical and chemical parameters that
eventually impact overall crop and soil viability (Visser and Parkinson, 1992).
Activity levels, biomass numbers, and community shifts can reflect the stability of
a system with respect to the level of nutrient cycling, the amount of carbon utilized
in a system, and the overall community structure and function in a soil system. Since
microbial diversity of agricultural soils is important to the maintenance of soil
formation, toxin removal, and elemental cycling (Borneman et al., 1996), diversity
measures will most likely become increasingly important in evaluations of soil
quality. Examples of rapid shifts in community structure such as these may serve
as early indicators of changes in soil quality (Turco et al., 1994).
Soil resilience is defined as the ability of the soil to recover after disturbance
(Elliott and Lynch, 1994). Soil resilience can be restored and biodiversity protected
by, among other things, reducing tillage and increasing crop rotations (Elliott and
Lynch, 1994). Studies have shown that by reducing the impact of disturbances, such
as tillage, microbial biomass levels and soil resilience will improve (Elliot and Lynch,
1994). In addition, crop rotations of at least 3 years can decrease disease-related
problems (Rovira et al., 1990), which can also increase crop resilience. However,
soil resilience may be different depending on the disturbance; disturbances such as
tillage may affect the general biological status, while pesticides or other stressors
may only affect individual functional groups (Swift, 1994). By further establishing
the relationships between the microbial biodiversity and soil resilience, a greater
understanding of microbial indicators can be achieved.
CONCLUSION
The diversity of microorganisms is greater than any other group of organisms

on Earth, but our knowledge of the diversity and genetic wealth in these groups is
limited. Microorganisms are key to the integrated functioning of nutrient cycles and
decomposition, soil structure, and plant growth in agricultural systems. Research is
needed to increase our understanding of the diversity and function of microbial
communities in agroecosystems. In agroecosystems, microbial diversity will influ-
ence all the other levels within the ecosystem by functions such as those involved
in nitrogen and carbon cycling, soil structure maintenance, and biological control.
The challenge ahead is to identify the level of microbial diversity, species compo-
sition and distribution, and the resiliency of the community to withstand stress and
maintain a quality ecosystem. We need to determine the extent of microbial diversity
in agroecosystems and increase our knowledge of the functional roles of microbes
to assess their role in agroecosystem quality and productivity.
ACKNOWLEDGMENTS
Support from the U.S. Department of Agriculture, Agricultural Research Service

in cooperation with the College of Agriculture and Home Economics, Washington

State University, Pullman, WA is appreciated. Trade names and company names are
included for the benefit of the reader and do not imply endorsement or preferential
treatment of the product by the U.S. Department of Agriculture or Washington State
University. All programs and services of the U.S. Department of Agriculture are
offered on a nondiscriminatory basis without regard to race, color, national origin,
religion, sex, age, marital status, or handicap.
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CHAPTER 2
Soil Microfauna: Diversity and

Applications of Protozoans in Soil
Stuart S. Bamforth
CONTENTS
Introduction
Role of Soil Protozoa
Measuring Protozoan Biodiversity
Protozoan Diversity in Agroecosystems
Applications
Conclusions
References
INTRODUCTION
Agricultural plant production depends upon the decomposition of plant and

animal residues, as well as fertilizers, into simpler compounds, many of which are
transformed into microbial and animal protoplasm. These organic materials are
eventually mineralized into simpler compounds, such as CO2, ammonia, and phos-
phate, which are absorbed by plant roots.
ROLE OF SOIL PROTOZOA
Microarthropods and larger fauna, especially earthworms, increase the rate and
amount of mineralization by comminution of organic matter and by redistribution
of “hot spots” of activity through movements. However, mineralization and return

of nutrients to plants occur in the water films covering soil aggregates and filling
their pores. Here, bacteria and fungi decompose organic matter and immobilize the
extracted nutrients into their bodies, but grazing by the microfauna, protozoa, and
nematodes regulates and modifies the size and composition of the microbial com-
munity and enhances microbial growth through microfaunal excretions. Nematodes
also graze fungi (Chapter 1), but protozoa, especially amoebae, can graze bacteria
in tiny pore spaces unavailable to nematodes. The degree of nutrient recycling is
influenced by external factors of climate and soil management (e.g., inputs of
fertilizers and biocides, compaction by farm machinery) and internally by the com-
munity of protozoa and nematodes, reflected in their biodiversity.
Most of the microfauna are located in small hot spots scattered through the soil
mosaic, which is soil aggregates of 1 mm or smaller, containing bits of organic
matter, detritus and the overlying litter, rhizosphere, and the “drillosphere” parts of
the soil influenced by earthworm secretions and castings. The microbial-feeding
microfauna constitute an essential component of the soil ecosystem; therefore,
changes in their community structure can influence mineralization and soil fertility.
MEASURING PROTOZOAN BIODIVERSITY
Soil protozoa comprise four groups: the “naked” rapidly growing flagellates,
amoebae, and ciliates, and the more slowly growing shelled amoebae, or testacea.
The small size and flexibility of the first two groups allows them to exploit small
pore spaces, and they furnish most of the protozoan numbers.
The more diverse and larger ciliates and testacea inhabit the larger pore spaces
which are subject to desiccation and other stresses; consequently, both groups show
a wide spectrum of species of r/K selection and degree of autochthonism (Wodarz
et al., 1992). Ciliates are divided into pioneer r-selected Colpodida, competitive K-
selected Polyhymenophora, and intermediate remaining taxa. Dividing the number
of species of the first group by the second produces a C/P ratio, where C/P > 1.00
indicates a stressed soil of low productivity, and C/P < 1.00 a more productive soil
with microarthropods and macrofauna (Foissner, 1987; Yeates et al., 1991). Among
the testacea, certain species indicate soil acidity or alkalinity, and the shell conveys
information about moisture fluctuations (Bonnet, 1964). Consequently, these two
groups can serve as bioindicators of soil conditions.
Ideally, biodiversity studies measure both species and numbers per species.
However, the small size and transparency of naked protozoa make them too difficult
to find among soil particles; consequently, counting has been traditionally performed
by the most probable number (MPN) technique of Singh (1946) or its modification
by Darbyshire et al. (1974). There are criticisms of the method (Foissner, 1987), in
response to which a second direct count method was developed by Griffiths and Ritz
(1988) that separates the protozoa from soil particles by percoll phosphate gradient
centrifugation and staining for fluorescent microscopy. This method is employed
routinely to measure the protozoan component of the soil fauna in experimental field
crop studies by the Technical University of Munich. The larger and more motile
ciliates can be counted by examining a watered soil suspension drop-by-drop until

at least 0.4 g of fresh soil has been examined (Foissner, 1987). The possession of a
shell enables direct counting of testacea by this method, or by staining a soil
suspension and mounting small samples on slides, providing a permanent record
(Couteaux, 1967; Korgonova and Geltser, 1977). Combining the temporary and
permanent methods provides a more complete census than either method alone.
Estimating species richness is best done by placing 10 to 50 g of sample in a
petri dish and adding water until 5 to 20 ml will drain off when gently pressed with
a finger. By placing several coverslips, each underlaid with a piece of lens paper on
top of the sample, and examining after 1 day, a variety of flagellate species will be
revealed. The culture is examined at 3 to 4 day intervals for a month to determine
the succession of species of mainly ciliates and testacea (Foissner, 1987). Most
amoebae will be found by streaking soil samples on bacterized water (non-nutrient)
agar plates or by placing soil samples in wells cut in such plates. The amoebae
migrate out from the soil particles (Bamforth, 1995a).
PROTOZOAN DIVERSITY IN AGROECOSYSTEMS
Studies on grasslands (McNaughton, 1977; Tilman, 1996) show that biodiversity

stabilizes community and ecosystem processes, although individual species within
the system may fluctuate considerably. Tilman (1996) found wide variations in the
biomass of the 24 most abundant species of plants in an 11-year study. In a 6-month
study of soil ciliates under a spruce stand, Lehle (1992) found that the proportions
of the three dominant ciliate species fluctuated widely: Cyclidium muscicola ranged
from 8 to 75% of the total populations and two colpodid species varied from 4 to
45%. The different responses in these two studies may reflect changes in the realized
niches of species; thus, biodiversity furnishes a reservoir of biotic abilities contrib-
uting to ecosystem sustainability (Bamforth, 1995b). Biodiversity, like the compar-
ison of nontillage to conventional agriculture, may not produce noticeable increases
in crop production, but maintaining biodiversity can retard the deterioration that has
characterized agroecosystems through 4000 years of human history. Protozoa can
serve as bioindicators of ecosystem conditions, and warn of soil impoverishment.
The appeal of protozoan bioindicators is their environmental sensitivity due to
their delicate cell membranes, their rapid growth rate, restricted movement in soil,
ubiquity, and wide range of morphologies in ciliates and testacea, providing a
multispecies approach enabling community analyses to indicate soil conditions
(Foissner, 1994). Difficulties arise in the taxonomy and time needed for identification
and enumeration, but, as the following applications illustrate, protozoa convey valu-
able information about agroecosystems because of their pivotal position in the
nutrient cycling that all terrestrial ecosystems depend upon.
APPLICATIONS
Conventional agriculture creates a special ecosystem by mixing the topsoil (and

compacting it) through tillage, removing plant canopies that protect the soil, adding

fertilizers and biocides, and removing harvests. A more sustainable agriculture
minimizes topsoil disturbance, reduces inputs, and substitutes organic for mineral
fertilizers (Doran and Werner, 1990).
Plowing in conventional agriculture incorporates crop residues into the soil
profile to produce homogeneous soils that favor the bacteria, protozoa, and bac-
tivorous nematode portions of the underground food web; in contrast, minimal
tillage leaves organic residues on the surface and a rich organic layer near the
surface, enhancing the fungal, Collembola, and earthworm portions of the under-
ground food web (Hendrix et al., 1986; Lee and Pankhurst, 1992). The protozoan
communities differ between the two systems in the greater prominence of r-selected
colpodid ciliates (reflecting less species diversity) in conventional fields (Foissner,
1992; Bamforth, unpublished data). The biomass of amoebae and flagellates, how-
ever, is greater in the surface layer of ecofarmed systems (DeRuiter et al., 1993)
and is associated with increased nitrogen mineralization (DeRuiter et al., 1993).
Using testacea as bioindicators, Wodarz et al. (1992) found organically farmed
field and vineyard soils showed improved soil conditions over conventionally
farmed counterparts.
Organic fertilizers, especially straw and animal manures, are more similar to
natural organic substrates than chemical fertilizers. Microbial and protozoan activity
is highest in organically enriched soils (Schnurer et al., 1985; Aescht and Foissner,
1991; 1992), and is usually accompanied by increases of most soil fauna, especially
earthworms (Doran and Werner, 1990), which enhance protozoan biodiversity.
The higher protozoan activity in soils under nontillage and organic fertilizer
management is enhanced by other fauna, especially earthworms, which disperse
bacteria and their protozoan predators to new locations, through burrowing move-
ments and passing ingested cysts through guts, providing new hot spots and releasing
greater quantities of nutrients, which have led to increased plant yields in a few
cases (Brown, 1995). Ingested active protozoa furnish a highly assimilable food
source, sustaining the fauna that enhance microbial and protozoan activities (Brown,
1995). Thus, high protozoan biodiversity usually reflects earthworm abundance.
The application of biocides often influences other organisms besides those tar-
geted. Herbicides have little effect on protozoa, although they may influence them
indirectly by altering bacterial nitrogen activities and by modifying the environment
in eliminating the vegetation over the soil. Insecticides and fungicides are more
toxic, as shown in the study of Petz and Foissner (1990) on the effects of lindane,
an insecticide, and mancozeb, a fungicide, on the soil ciliate and testacean commu-
nities of a spruce forest. The insecticide decreased both numbers and species, and
altered the community structure of ciliates by increasing the abundance of several
colpodids. This result shows the value of multispecies-monitoring studies, and also
the value of biodiversity to an ecosystem, allowing response to changing conditions
(Bamforth, 1995b). The insecticide exerted less effect on testacea, and the fungicide
exerted little influence on both groups. The investigation used a randomized block
design and extended the study period to the 90 days needed to ascertain if the biocide
caused acute toxicity (Domsch et al., 1983). This type of study shows the precision
that protozoan bioindicators can provide to assess agroecosystem conditions.

The heavy machinery used in modern farming causes soil compaction, destroying
not only the worm and root channels that reduce soil porosity and the larger fauna,
but also reducing pore spaces in which bacteria and their protozoan predators live.
Compaction reduces testacean species diversity and eliminates large forms (Berger
et al., 1985), and a number of studies relating pore space to protozoan activity (e.g.,
Rutherford and Juma, 1992; England et al., 1993 ) show less activity in smaller
spaces. Griffiths and Young (1994) found the same trend and concluded that com-
paction influences protozoa indirectly by producing anaerobic conditions that inhibit
protozoa and reduce the metabolism and reproduction of their bacterial prey.
A vital part of agricultural management is soil conservation and restoration,
which can be monitored by analyzing the protozoan community to assess the degree
of the comprehensive biological activity to productive farming (Yeates et al., 1991;
Wodarz et al., 1992).
CONCLUSIONS
Protozoa and nematodes are pivotal organisms in agroecosystems because their

predation upon bacteria increases mineralization of nutrients necessary for plant
growth. Since biodiversity stabilizes community and ecosystem processes (Tilman,
1996), maintaining and increasing protozoan biodiversity will contribute to more
sustainable agriculture. Ecofarming and organic fertilizer management enhance pro-
tozoan activity.
Protozoa have several unique features, such as rapid sensitivity to environmental
changes and ubiquity, that favor their use as bioindicators. Protozoan biodiversity
reflects the condition of an agroecosystem and can be used to monitor the effects
of environmental changes.
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CHAPTER 3
Diversity and Function of Soil Mesofauna
Deborah A. Neher and Mary E. Barbercheck
CONTENTS
Introduction
Habitat
Biology and Ecology of Soil Fauna
Plant Feeders
Microbial Feeders
Omnivory
Predators
Ecosystem Processes
Value of Diversity
Agricultural Disturbances
Soil Texture and Compaction
Cultivation
Fertilization
Pesticides
Concluding Remarks
References
INTRODUCTION
Diversity in natural communities of microbes, plants, and animals is a key factor

in ecosystem structure and function. Agricultural ecosystems, however, are designed
around one or several species of plants or animals. Reduction of diversity in agri-
cultural systems, compared with that in natural ecosystems, is traditionally consid-
ered essential to increase production of food, forage, and fiber. For simplicity of

Table 1 Hierarchy of Size and Abundance of Organisms Inhabiting Soil
Biomass Length Populations
Class Example(s) (g m–2) (mm) (m–2)
Microflora Bacteria, fungi, algae, 1–100 Not applicable 106–1012
actinomycetes
Microfauna Protozoa 1.5–6.0 0.005–0.2 106–1012
Mesofauna Nematodes, 0.01–10 0.2–10 102–107
enchytraeids,
mites, Collembola
Macrofauna Insects 0.1–2.5 10–20 102–105
Megafauna Earthworms 10–40 20 0–103
Data from Dindal (1990) and Lal (1991).
management, biological cycles are sometimes replaced by fossil fuel-based products,

e.g., synthetic fertilizers. Intense management practices that include application of
pesticides and frequent cultivation affect soil organisms, often altering community
composition of soil fauna. Soil biological and physical properties (e.g., temperature,
pH, and water-holding characteristics) and microhabitat are altered when native
habitat is converted to agricultural production (Crossley et al., 1992). Changes in
these soil properties may be reflected in the distribution and diversity of soil meso-
fauna. Organisms adapted to high levels of physical disturbance become dominant
within agricultural communities, thereby reducing richness and diversity of soil
fauna (Paoletti et al., 1993).
Relationships between particular groups of organisms and management practices
in agriculture can be studied under specific circumstances to define expected levels
of diversity. The full diversity of soil communities has not been quantified for either
agricultural or native ecosystems (Lee, 1991), and, in addition, the relationship
between biodiversity and ecosystem function is not understood fully (Walker, 1992).
Theoretically, this knowledge could be used to establish and maintain conditions
that optimize beneficial effects of these organisms. Realistically, however, ideal
conditions may be difficult to attain because of constraints imposed by agricultural
production practices. We do not have sufficient knowledge to determine whether or
not it is necessary, possible, or desirable to duplicate in agriculture the biodiversity
that may be present in natural ecosystems.
This chapter examines the diversity and some of the functions of soil mesofauna
in agricultural systems (Table 1). Most research on soil biota has focused on eco-
systems such as forests and grasslands that are managed less intensively than agri-
cultural or row crop systems. Ecologists have paid more attention to the role of
micro- and mesofauna in ecosystem function, whereas agricultural scientists have
focused on their role in nitrogen fixation and as pests and pathogens of crops. Our
understanding of the role of soil organisms in agricultural systems is increasing, but
more research is needed to elucidate their significance to crop production. Mesofauna
occupy all trophic levels within the soil food web and affect primary production
directly by root feeding and indirectly through their contribution to decomposition
and nutrient mineralization (Crossley et al., 1992). Detailed reviews of the biology
of soil fauna and their relationship to soil structure and ecological function are

available (Wallwork, 1976; Swift et al., 1979; Freckman, 1982; Peterson and Luxton,
1982; Pimm, 1982; Seastedt, 1984; Dindal, 1990; Beare et al., 1992).
HABITAT
Unlike soil macrofauna (e.g., earthworms, termites, ants, some insect larvae),
mesofauna generally do not have the ability to reshape the soil and, therefore, are
forced to use existing pore spaces, cavities, or channels for locomotion within soil.
Habitable pore space (voids of sufficient size and connectivity to support mesofauna)
accounts for a small portion of total pore space (Hassink et al., 1993b). Microfaunal
community composition becomes increasingly dominated by smaller animals as aver-
age pore volume decreases. Within the habitable pore space, microbial and mesofaunal
activity is influenced by the balance between water and air. Maximum aerobic micro-
bial activity occurs when 60% of the pore volume is filled with water (Linn and Doran,
1984). Saturation (waterlogging) and drought are detrimental to soil faunal commu-
nities because these conditions result in anaerobiosis or dehydration, respectively.
Populations and diversity of mesofauna are greatest in soil with high porosity
and organic matter, and structured horizons (Andrén and Lagerlöf, 1983). Most
biological activity occurs within the top 20 cm of soil which corresponds to the
“plow layer” in agricultural soils. In uncultivated soil, mesofauna are more abundant
in the top 5 cm than at greater depths in the soil. The organic horizon (O) is the
area of accumulation of recognizable plant materials (high C:N ratio) and animal
residues (low C:N ratio). The fermentation (F or O1) layer consists of partially
decomposed, mixed plant and animal debris permeated with hyphae of fungi and
actinomycetes. The humus (H or O2) horizon contains amorphous products of decom-
position with the source unrecognizable. Eventually, organic matter from these
horizons becomes incorporated into the mineral soil profile. Because cultivated
agricultural systems often lack a distinct organic layer on the surface, one might
expect diversity of soil biota to be less than in uncultivated or no-till soils (House
et al., 1984).
Plants affect soil biota directly by generating inputs of organic matter above-
and belowground and indirectly by the physical effects of shading, soil protection,
and water and nutrient uptake by roots. Energy and nutrients obtained by plants
eventually become incorporated in detritus which provide the resource base of a
complex soil food web. Plant roots also exude amino acids and sugars which serve
as a food source for microorganisms (Curl and Truelove, 1986). Soil mesofauna are
often aggregated spatially which is probably indicative of the distribution of favored
resources, such as plant roots and organic debris (Swift et al., 1979; Goodell and
Ferris, 1980; Barker and Campbell, 1981; Noe and Campbell, 1985).
BIOLOGY AND ECOLOGY OF SOIL FAUNA
Soil mesofauna are often categorized by specific feeding behaviors, often depicted
as microbial feeders. However, it should be emphasized that many organisms are at

least capable of feeding at other trophic groups. As a result, omnivory in soil
communities may be more prevalent than assumed previously (Walter et al., 1986;
Walter, 1987; Walter et al., 1988; Walter and Ikonen, 1989; Mueller et al., 1990).
Our discussion will focus specifically on nematodes, Collembola (springtails), and
mites because they predominate in total numbers, biomass, and species of fauna in
soil (Harding and Studdart, 1974; Samways, 1992).
Soil nematodes are relatively abundant (6 × 104 to 9 × 106 per m2), small (300
µm to 4 mm) animals with short generation times (days to a few weeks) that allow
them to respond to changes in food supply (Wasilewska, 1979; Bongers, 1990).
Relative to other soil microfauna, trophic or functional groups of nematodes can be
identified easily, primarily by morphological structures associated with various
modes of feeding (Yeates and Coleman, 1982; Freckman, 1988; Bongers, 1990).
Nematodes may feed on plant roots, bacteria, fungi, algae, and/or other nematodes
(Wasilewska, 1979).
Mites and collembolans can account for 95% of total soil microarthropod num-
bers (Harding and Studdart, 1974). Soil mites occur mainly in three suborders. The
suborder Oribatida (Cryptostigmata) comprises the numerically dominant group in
the organic horizons of the soil. Members of the mite suborder Mesostigmata
(Gamasida) are relatively large, active mites. The mite suborder Prostigmata (Actine-
dida) is a large and taxonomically complex group. Soil prostigmatids have more
heterogeneous feeding habits than other mite suborders (see table in Kethley, 1990,
for feeding habits). Prostigmatids are mostly fungal feeders and predators.
Collembolans are abundant and distributed widely. Collembolans have relatively
high metabolic, feeding, and reproductive potential. Functional classification of
collembolans (Christiansen, 1964; Bödvarsson, 1970; Verhoef and Brussard, 1990)
can be based on gut content or shape of the mouthparts, which are adapted to the
specific feeding habit (Swift et al., 1979). Because most forms of Collembola feed
on decaying vegetation and associated microflora, the distribution of mycelia and
spores of saprophytic fungi may be a major factor influencing the distribution of
collembolans.
Other groups of arthropods that occur commonly in soil are pseudoscorpions,
symphylans, pauropods, proturans, diplurans, and the immature stages of holome-
tabolous insects (Dindal, 1990). Ants and termites can also be very numerous;
however, these macroarthropods will not be considered here (Brian, 1978).
Plant Feeders
Plant-feeding nematodes can become abundant in agricultural ecosystems

(Wasilewska, 1979; Popovici, 1984; Neher and Campbell, 1996). These nematodes
may affect primary productivity of plants by altering uptake of water and nutrients.
These abnormalities may result from changes in root morphology and/or physiology.
For many agricultural crops, a negative relationship between crop yield and popu-
lations of plant-feeding nematodes, such as Meloidogyne, Heterodera, and Praty-
lenchus spp., has been observed (Mai, 1985; Barker et al., 1994). However, when
entire nematode communities, including free-living nematodes, are examined, a

positive association has been observed between plant biomass production and total
nematode populations in grassland ecosystems (Yeates and Coleman, 1982). This
relationship holds for plant production measured as harvested hay and root biomass
(King and Hutchinson, 1976). A negative relationship between total nematode pop-
ulations and plant productivity has been observed in tropical forests (Kitazawa,
1971). The relationship between soil nematode communities and row crop yield has
yet to be determined.
Microarthropods rarely harm crop plants. However, soil mesofauna may
become pests when a preferred food source is absent. Some Collembola, e.g.,
sminthurids and onychiurids, may feed on roots. For example, root-grazing injury
on sugar beet is caused by Onychiurus spp. (Collembola) rubbing their bristled
bodies against roots (Curl et al., 1988). However, root injury decreases if specific
weed species and certain kinds and amounts of organic matter are present and,
thus, provide the preferred microbial food supply. Few groups of soil mites are
adapted to feeding on live plant tissues in soil. Some examples occur in the
Tarsonemidae (Prostigmata) and Periohmanniidae (Oribatida). Most soil mites feed
on plant material only after decomposition has begun. Often, increasing vegeta-
tional diversity and the quality and quantity of organic matter in soil increases
potential benefits by soil mesofauna.
Microbial Feeders
Microbial-feeding mesofauna feed on fungi (including mycorrhizae), algae,

slime molds, and bacteria by removing them from clumps of decaying material or
soil aggregates (Moore and de Ruiter, 1991). Generally, bacterial-feeding nematodes
such as Cephalobidae and Rhabditidae (Neher and Campbell, 1996) are abundant
in agricultural ecosystems (Wasilewska, 1979; Popovici, 1984). Consumption of
microbes by soil mesofauna alters nutrient availability by stimulating new microbial
growth and activity plus releasing nutrients immobilized previously by microbes.
In general, fungal feeding is the dominant trophic function of microarthropods.
Collembolan species have preferred food sources which are maintained even after
the material has passed through the digestive tracts of other animals. For example,
the collembolans Proisotoma minuta and O. encarpatus feed upon the soilborne
fungal plant pathogen Rhizoctonia solani which causes damping-off disease on
cotton seedlings (Curl et al., 1988). These collembolan species prefer feeding on
the fungal pathogen in soil compared with the biocontrol fungi Laetisaria arvalis,
Trichoderma harzianum, and Gliocladium virens (Curl et al., 1988). Additionally,
collembolan species can distinguish and graze selectively on different species of
vesicular-arbuscular mycorrhizae (Thimm and Larink, 1995).
Almost all oribatid mites are microbial feeders. Examples of microbial feeding
also occur in the Mesostigmata (Ameroseiidae, Uropodidae) and Prostigmatida
(Tarsonemidae, Nanorchestidae, Stigmaeidae Pygmephoridae, Eupodidae, and
Tydeida). Although many microarthropods are microbial feeders, recent studies
indicate that other arthropods are omnivorous and shift feeding behavior as food
resources change (Walter, 1987; Mueller et al., 1990).

Omnivory
Omnivores add “connectedness” to the food web by feeding on more than one
food source (Coleman et al., 1983). Omnivorous nematodes, such as some Doryla-
midae, make up only a small portion of the total nematodes in agricultural ecosystems
(Wasilewska, 1979; Neher and Campbell, 1996). They may feed on algae, bacteria,
fungi, and other nematodes. Collembolans are often microbial feeders, but may also
be facultative predators of nematodes (Snider et al., 1990). Mites that feed on both
microbes and decaying plant material can be found in the oribatid mite families
Nothridae, Camisiidae, Liacaridae, Oribatulidae, and Galumnidae. Coprophages,
which ingest dung and carrion, including dead insects, are found among the oribatid
families Euphthiracaridae, Phthiracaridae, Galumnidae, and Oppiidae.
Predators
Mesofauna may be predators or serve as prey for predaceous mites and other
predators, such as beetles, fly larvae, centipedes, and spiders. Predatory nematodes
feed upon all the other trophic groups of nematodes (Moore and de Ruiter, 1991)
and represent only a small portion of the total nematodes in agricultural ecosystems
(Wasilewska, 1979). Nematode predators (e.g., members of the orders Mononchida
and Tripylida) and insect-parasitic nematodes (e.g., members of the families Stein-
ernematidae, Diplogasteridae, Mermithidae) present in the soil may affect popula-
tions of their prey (Poinar, 1979; Small, 1987; Stirling, 1991).
Soil microarthropods can be important predators on small arthropods (e.g.,
proturans, pauropods, enchytraeids) and their eggs, nematodes, and on each other.
Predation of insect eggs in agroecosystems may constitute a major influence of
controlling microarthropod populations. Brust and House (1988) found that the mite
Tyrophagus putrescentiae is an important predator of eggs of southern corn rootworm
Diabrotica undecimpunctata howardi in peanuts. Chaing (1970) estimated that pre-
dation by mites accounted for 20% control of corn rootworms (Diabrotica spp.) and
63% control following the application of manure. Mite predation on root-feeding
nematodes may be significant under some conditions (Inserra and Davis, 1983;
Walter, 1988). For example, one adult of the mesostigmatid mite Lasioseius scap-
ulatus and its progeny consumed approximately 20,000 Aphelenchus avenae on agar
plates in 10 days (Imbriani and Mankau, 1983). Collembolan species may also
consume large numbers of nematodes (Gilmore, 1970). For example, Entomobry-
oides dissimilis consumed more than 1000 nematodes in a 24-h period. Furthermore,
collembolans may consume large numbers of insect-parasitic nematodes and, thus,
affect the efficacy of these nematodes used as biological control agents of soil-
dwelling insect pests (Epsky et al., 1988; Gilmore and Potter, 1993).
ECOSYSTEM PROCESSES
Micro- and mesofauna contribute directly to ecosystem processes such as decom-

position and nutrient cycling in complex and interactive ways (Swift et al., 1979).

Bacteria, actinomycetes, fungi, algae, and protozoa are primary decomposers of
organic matter. These microorganisms are involved directly with production of
humus, cycling of nutrients and energy, elemental fixation, metabolic activity in soil,
and the production of complex chemical compounds that cause soil aggregation.
Microbial-grazing mesofauna affect growth and metabolic activities of microbes
and alter the microbial community, thus regulating decomposition rate (Wasilewska
et al., 1975; Trofymow and Coleman, 1982; Whitford et al., 1982; Yeates and
Coleman, 1982; Seastedt, 1984) and nitrogen mineralization (Seastedt et al., 1988;
Sohlenius et al., 1988). Nematodes feed on bacteria and fungi on decaying organic
matter, but not on the organic matter itself. Nematode species with a buccal stylet
(spearlike structure) feed on cell contents and juices obtained by piercing the cellular
walls of plant roots or fungal mycelium. Other species have no stylets and feed on
particulate food such as bacteria and small algae (Vinciguerra, 1979). Microarthro-
pods fragment detritus and increase surface area for further microbial attack (Berg
and Pawluk, 1984). For example, collembolans and mites may enhance microbial
activity, accelerate decomposition, and mediate transport processes in the soil. Even
when they do not transform ingested material significantly, they break it down,
moisten it, and make it available for microorganisms.
There is evidence that plants benefit from increased mineralization of nitrogen
by soil mesofauna. Shoot biomass and nitrogen content of plant shoots grown in the
presence of protozoans and nematodes were greater when compared with plants
grown without mesofauna (Verhoef and Brussard, 1990). Soil fauna are responsible
for approximately 30% of nitrogen mineralization in agricultural and natural eco-
system soils. The main consumers of bacteria are protozoa and bacterial-feeding
nematodes which account for 83% of nitrogen mineralization contributed by soil
fauna (Elliott et al., 1988). Nematodes also excrete nitrogenous wastes, mostly as
ammonium ions (Anderson et al., 1983; Ingham et al., 1985; Hunt et al., 1987).
Collembola excrete nitrate in concentrations 40 times more than their food source
(Teuben and Verhoef, 1992). Furthermore, large collembolan species increase min-
eralization by selective feeding on fungi, whereas smaller species aid in the formation
of humus by nonselective scavenging and mixing of the mineral and organic fractions
of soil (van Amelsvoort et al., 1988). Microfauna constitute a reservoir of nutrients.
When microfauna die, nutrients immobilized in their tissues are mineralized and
subsequently become available to plants.
Soil fauna transport bacteria, fungi, and protozoa (in gut or on cuticle) across
regions of soil and, thus, enhance microbial colonization of organic matter (Seastedt,
1984; Moore et al., 1988). For example, Collembola and sciarid fly larvae transmit
root-infecting fungi and fungal parasites (Anas and Reeleder, 1988; Whipps and
Budge, 1993). Microarthropods are surrounded by and, therefore, may disseminate
propagules of insect-parasitic fungi including Beauveria spp., Metarhizium spp.,
Paecilomyces spp., and Verticillium spp. and facultative pathogens of insects in the
genera Aspergillus and Fusarium spp. Under laboratory conditions, Collembola and
mites transport spores of the insect-parasite M. anisopliae (Zimmerman and Bode,
1983). The impact of insect-parasitic fungi on natural populations of microarthropods
is unknown.

VALUE OF DIVERSITY
Diversity in form and function of biotic communities results in the formation of

spatial and temporal heterogeneity of organisms that contributes to the overall function
of the ecosystem. Individual taxa may have multiple functions, and several taxa may
appear to have similar functions. However, function may not necessarily be redundant,
because taxa performing the same function are often isolated spatially, temporally, or
by microhabitat preference (Beare et al., 1995). Biodiversity allows organisms to avoid
intense competition for food or space, decrease invasion and disruption, and maintain
constancy of function through fluctuating environmental conditions.
Various measures of diversity are available to describe soil invertebrate commu-
nities including abundance, biomass, density, species richness, species evenness,
maturity indexes, trophic/guild structure, and food web structure. Indexes of diversity,
which include elements of richness (number of taxa) and evenness (relative abun-
dances), can be applied at scales ranging from alleles and species to regions and
landscapes. Diversity indexes do not reveal the taxonomic composition of the com-
munity. For example, a community composed entirely of exotic species could have
the same index value as a community composed entirely of endemic species. There-
fore, a diversity index, by itself, does not predict ecosystem health or productivity.
Debates concerning relationships between biodiversity and ecosystem stability
became popular in the 1960s and 1970s. MacArthur (1955) was the first to argue
that complex systems are more stable than simple systems. In the early 1970s, May
(1972; 1973) used mathematical models to argue that diverse communities were less
stable than simple systems. Today, some conclude that relatively simple, short food
webs that exhibit little omnivory or looping are more stable than longer food webs
with much omnivory or looping. A short food web is one with few trophic levels
(Polis, 1991). Others hypothesize that high linkage is responsible for making food
webs unstable, i.e., stability can develop if numbers of species increase but not if
omnivory increases (Pimm et al., 1991; Lawton and Brown, 1993). It is clear from
this ongoing debate that it is impossible to generalize the relationship between
biodiversity and ecosystem stability. Besides, none of the theories has been tested
adequately for application to soil communities.
Factors affecting diversity within trophic groups of the detritus food web include
altitude, latitude (Procter, 1984; Rohde, 1992), predation in the presence of strong
competitive interactions (Petraitis et al., 1989), and disturbance (Petraitis et al., 1989;
Hobbs and Huenneke, 1992). For example, the pervading theory is that the greatest
species diversity is found in the tropics and that diversity decreases with increasing
latitude (Rohde, 1992). However, the opposite is true for free-living nematodes.
Free-living nematodes are more diverse and abundant in temperate than in tropical
regions (Procter, 1984; 1990). Nematodes are tolerant of harsh conditions at high
latitudes but are not competitive against more-specialized soil fauna in the tropics
(Petraitis et al., 1989).
At smaller scales, predators may promote species diversity among competing
prey species when they feed preferentially on exceptionally competitive prey (Petrai-
tis et al., 1989). Disturbance also plays a role with the “intermediate disturbance

hypothesis” suggesting that taxonomic diversity should be highest at moderate levels
of disturbance (Petraitis et al., 1989; Hobbs and Huenneke, 1992). Disturbance is
defined as a cause (a physical force, agent, or process, either abiotic or biotic) that
results in a perturbation (an effect or change in system state relative to a reference
state and system) (Rykiel, 1985). If disturbance is too mild or too rare, then soil
communities will approach equilibrium and will be dominated by fewer taxa that
can outcompete all other taxa. However, attainment of steady-state equilibria in
agricultural or natural ecosystems is uncommon (Richards, 1987). If disturbance is
common or harsh, only a few taxa that are insensitive to disruption will persist,
therefore decreasing biodiversity (Petraitis et al., 1989). For example, Prostigmatid
mites in the Eupodidae, Tarsonemidae, and Tydeidae are among the most abundant
in cultivated agroecosystems and their numbers increase rapidly in response to
disturbances such as cultivation (Crossley et al., 1992).
AGRICULTURAL DISTURBANCES
Disturbance can alter the diversity of an ecosystem (Atlas, 1984) directly by

affecting survivorship of individuals or indirectly by changing resource levels (Hobbs
and Huenneke, 1992). Sometimes, diversity measurements reflect the result of dis-
turbance caused by pollution and/or stress. For example, taxonomic diversity of
microinvertebrate communities was less in polluted or disturbed than in unpolluted
or undisturbed agricultural sites (Atlas et al., 1991). Pollution eliminates sensitive
species, reducing competition so that tolerant species proliferate (Atlas, 1984).
The successional status of a soil community may also reflect the history of
disturbance. Succession in cropped agricultural fields begins with depauperate soil
which acts like an island to which a series of organisms immigrate. First, opportu-
nistic species, such as bacteria and their predators, are colonists of soil. Subsequently,
fungi and their predators migrate into the area (Böstrom and Sohlenius, 1986).
Microarthropods, such as collembolans, mites, and fly maggots, can colonize nearly
bare ground and rise quickly in population density. Top predator microarthropods,
such as predaceous mites and nematodes, become established later and may have a
function similar to keystone predators in other community food webs (Elliott et al.,
1988). Finally, macro- and megafauna, such as earthworms, millipedes, slugs, cen-
tipedes, wood lice, sow bugs, and pill bugs, join the soil community (Strueve-
Kusenberg, 1982).
Succession can be interrupted at various stages by agricultural practices, such
as cultivation and applications of fertilizer and pesticide (Ferris and Ferris, 1974;
Wasilewska, 1979). Such interruptions reduce diversity and successional “maturity.”
Maturity indices are based on the principles of succession and relative sensitivity of
various nematode taxa to stress or disruption of the successional sequence (Bongers,
1990). Indices that describe associations within biological communities, such as a
maturity index, are less variable than measures of abundance of a single taxonomic
or functional group and are, thus, more reliable as measures of ecosystem condition
(Neher et al., 1995).

Soil Texture and Compaction
Soil texture may impose physical restrictions on the ability of fauna to graze on
microbes; therefore, texture may play a role in faunal-induced mineralization of
microbial carbon and nitrogen (van Veen and Kuikman, 1990). Carbon and nitrogen
mineralization is generally faster in coarse than in fine-textured soils. In clay soils,
organic material is protected physically from decomposers by its location in small
pores. In sandy soils, organic matter is protected by its association with clay particles
(Hassink et al., 1993a). Nematodes and microarthropods are often less abundant in
heavy clay soil than in sandy or peat soil (van de Bund, 1970; Zirakparvar et al.,
1980; Verma and Singh, 1989). Euedaphic species such as collembolans in the
Onychiuridae and mesostigmatid mite Rhodacarus roseus are especially rare in clay
soil (Didden, 1987).
Mesofauna are affected adversely by soil compaction (Aritajat et al., 1977a,b).
Wheel-induced compaction reduces soil porosity, which is accompanied by a
decrease in microbial biomass carbon and the density of Collembola (Heisler and
Kaiser, 1995). Collembolans avoid narrow pores to protect their waxy surface from
damage (Choudhuri, 1961). Wheel traffic decreased the density of collembolans and
predatory mites by 30 and 60%, respectively, compared with noncompacted soil.
The number of species was also reduced by compaction (Heisler, 1994).
Cultivation
Cultivation affects biogeochemical cycling by physically rearranging soil parti-

cles and changing pore size distribution, patterns of water and gas infiltration, and
gas emission (Klute, 1982). Tillage disrupts soil aggregates, closes soil cracks and
pores, and promotes drying of the surface soil. Soil fauna become sparse in top
layers of cultivated soil because moisture content fluctuates widely and the original
pore space network in this layer is destroyed. These physical alterations of the surface
layers of soil may persist for many years after cultivation has ceased.
Soils managed by conventional — or reduced — tillage practices have distinct
biological and functional properties (Doran, 1980; Hendrix et al., 1986). Plant
residue is distributed throughout the plow layer in fields managed with conventional
tillage. Under these conditions enhanced by cultivation, organisms with short gen-
eration times, small body size, rapid dispersal, and generalist feeding habits thrive
(Steen, 1983). These soils are dominated by bacteria and their predators such as
nematodes and astigmatid mites (Andrén and Lagerlöf, 1983; Yeates, 1984; Hendrix
et al., 1986; Beare et al., 1992) and are considered in an early stage of succession.
Oribatid and mesostigmatid mites decrease while other groups such as prostigmatid
mites and Collembola tolerate, but do not benefit, from cultivation (Crossley et al.,
1992). However, prostigmatid mite communities can be more diverse, containing
both fungal- and nematode-feeding taxa in cultivated soils (van de Bund, 1970).
Many microarthropods have omnivorous feeding habits in systems cultivated fre-
quently (Beare et al., 1992).
Conservation — or no-till — practices generate more biologically complex soils
than conventional tillage; however, in general, no-tillage cultivation does not appear

to result in greater concentrations of microarthropods than conventional tillage
except under drought stress (Perdue and Crossley, 1989). However, many studies
comparing tillage effects are short term. Our knowledge about tillage effects may
change as more long-term studies are implemented. Reduced tillage leaves most of
the residue of the previous crops on the soil surface, and results in changes in physical
and chemical properties of the soil (Blevins et al., 1983). Surface residues retain
moisture, dampen temperature fluctuations, and provide a continuous substrate
which promotes fungal growth. The increased fungal abundance can be attributed
to the ability of fungi to translocate nutrients from soil into surface residues, their
tolerance of lower pH and water potentials that often occur in surface residues, and
their ability to penetrate and use large detritus particles (Hendrix et al., 1986; Holland
and Coleman, 1987). Relative abundances of fungi and their predators, such as
nematodes and many microarthropods groups (e.g., uropodid mesostigmatid mites;
tarsonemid, eupodid, tydeids, and pygmephorid prostigmatid mites; oribatid mites)
(Walter, 1987), in no-till soils represent a more mature successional state than one
dominated by bacteria (Yeates, 1984; Böstrom and Sohlenius, 1986; Hendrix et al.,
1986; Holland and Coleman, 1987; Neher and Campbell, 1994). Fungal feeding by
microarthropods may stimulate microbial growth and enhance decomposition and
nutrient immobilization (Seastedt, 1984). However, nutrient mineralization rates are
relatively slow with stratification of debris and soil; nutrients are immobilized inside
plant debris on the soil surface (Hendrix et al., 1986; Holland and Coleman, 1987).
Fertilization
Fertilization may influence the population abundance or composition of meso-

faunal communities in soil. The outcome is a result of factors such as fertilizer
quality and/or quantity (Verhoef and Brussard, 1990). Fertilization may, thus, affect
the abundance and diversity of soil mesofauna directly or indirectly. These changes
in community composition may, in turn, influence ecosystem function.
Nutrients applied to agricultural soils may be derived either from fossil fuels or
plant and animal waste products. Nutrients are available in both forms, but organic
amendments also contain microbes and their respective food resources. Additions
of mineral fertilizers decrease populations of oribatid (cryptostigmatid) and prostig-
matid mites, and root- and fungal-feeding, omnivorous and predaceous nematodes
(Sohlenius and Wasilewska, 1984). Numbers of root-feeding nematodes may
increase with increased nitrogen fertilizer (Wasilewska, 1989). Populations of astig-
matid mites (Andrén and Lagerlöf, 1983) and bacterial-feeding nematodes (Sohle-
nius and Wasilewska, 1984) increase with additions of mineral fertilizer, but even
more so when soils are fertilized with manure which simultaneously adds organic
matter and microbes (Andrén and Lagerlöf, 1983; Weiss and Larink, 1991). However,
in Dutch polder soil, abundances and biomass of nematodes, mites, and Collembola
were similar between fields fertilized with manure, crop residues, and green manure
and those field soils amended with crop residues and synthetic fertilizer (van de
Bund, 1970).
Mesofauna aggregate around manure and plant litter (van de Bund, 1970). Pop-
ulations of fungal-feeding nematodes (Weiss and Larink, 1991), potworms

(Enchytraeids), collembolans, and sometimes mesostigmatid mites increase with
applications of manure (Andrén and Lagerlöf, 1983). Fratello et al. (1989) examined
the effects of seven types of organic fertilizer on the microfauna in alfalfa fields.
The reactions of populations of microarthropods to the different treatments varied
with sample date, illustrating the highly complex interactions that occur in the soil.
Poultry manure, sheep manure, worm compost, autoclaved urban sludge, urban
sludge, vetch green manure, or straw were added to soil to provide a common level
of 4% organic matter. Straw was the only additive that did not depress mite popu-
lations. Fewer mites and Collembola were found in plots treated with autoclaved
urban sludge than those treated with non-autoclaved urban sludge.
The quality of plant and animal wastes as nutrient sources may be altered by
composting. For example, applications of aged compost can increase suppression
of plant pathogens by increasing the effectiveness of biocontrol agents. The plant-
pathogenic fungus Rhizoctonia solani may cause damping-off disease in soil when
fresh or immature compost material high in cellulose content is added. However, in
aged compost, cellulose is degraded and the biocontrol fungus Trichoderma spp.
can parasitize the pathogen effectively, thus suppressing disease (Chung et al., 1988).
Large doses of mineral or manure fertilizers can harm mesofauna because of
toxicity (e.g., anhydrous ammonia) or high osmotic pressure due to salt (Andrén
and Lagerlöf, 1983). The repellent nature of ammonium can affect soil invertebrates
adversely (Potter, 1993). The potential for toxic effects can be decreased by applying
composted manure and sludge (Ott et al., 1983). However, accumulation of heavy
metals from repetitive sludge applications may kill omnivorous and predaceous
nematodes (Weiss and Larink, 1991).
Fertilization affects soil microflora and, thus, indirectly impacts soil mesofauna
by changing their food resources (Weil and Kroontje, 1979). Additions of nitrogen
may acidify soil and inhibit microbial growth and activity. Nitrogen may also affect
the quality of microbes as a food source for mesofuana. Booth and Anderson (1979)
grew two species of fungi in liquid media with 2, 20, 200, or 2000 ppm nitrogen
and determined the fecundity of the collembolan Folsomia candida while feeding
on the fungi. Fecundity increased with increasing nitrogen content up to 200 ppm.
F. candida did not show a preference for feeding on fungi with a greater or lesser
nitrogen content.
The effect of fertilization on microarthropod species diversity and abundance
within taxa and the subsequent impact on decomposition and nutrient mineralization
processes are not well understood. For example, synthetic fertilizers increase nem-
atode diversity, but applications of manure decrease nematode diversity (Wasilewska,
1989). The mechanism(s) explaining the differences are not understood. Applications
of synthetic nitrogen fertilizer on Swedish arable soils growing spring barley (Hor-
deum distichum L.) changed community composition, but not numbers and biomass
of nematodes, Collembola and mites (Andrén et al., 1988). Within a given environ-
ment, increased densities of microarthropods have been correlated with increased
foliage, root and microbial productivity (Lussenhop, 1981), or increased food
resource via fertilization. It is not known at what scale of resolution soil faunal
communities respond to changes in ecosystem function.

Pesticides
Pesticides are an integral part of modern farming practice. Pesticides can enter
the soil by a variety of routes, e.g., intentional application, spillage, overspraying,
runoff, aerial transport with soil, or leaching. Organic matter plays a major role in
the binding of pesticides in soil. Fulvic and humic acids are most commonly involved
in binding interactions. Pesticides or their degradation intermediates can also be
polymerized or incorporated into humus by the action of soil microbial enzymes
(Bollag et al., 1992).
Soil fumigation with general biocides such as methyl bromide decreases micro-
bial populations and nearly eliminates nematodes (Yeates et al., 1991). Although
recovery occurs, population densities may not return to prefumigation levels even
after 5 months (Yeates et al., 1991). Fumigation with general biocides return the
successional status of soil to that of a depauperate soil matrix that can only be
inhabited by primary colonists. However, within 60 weeks after soil fumigation and
manuring, a progression of colonization by early successional species followed by
more-specialized, later successional taxa can be observed (Ettema and Bongers,
1993).
Broad-spectrum insecticides that are applied for the control of insect pests can
be toxic to predaceous and parasitic arthropods. A single surface application of
chlorpyrifos reduced populations of predatory mites in plots of Kentucky bluegrass
for 6 weeks and similar applications of isofenphos reduced populations of non-
oribatid mites, Collembola, millipedes, and Diplura for as long as 43 weeks (Potter,
1993). Densities of Collembola were lower in aldicarb-treated soil than in untreated
soil, but only the collembolans in the suborder Arthropleona were influenced nega-
tively, whereas Symphypleona were not affected or occurred in higher numbers in
soil treated with aldicarb (Koehler, 1992). Mesostigmatid mites did not occur at the
site for the first 2 months after treatment, and their abundance was reduced for 6
months. After 3 and 4 years, abundance was similar in treated and untreated soil.
Koehler (1992) noted a change in species composition associated with aldicarb
treatment and categorized three groups of reaction. The most sensitive organisms
were absent from 9 months to 1 year after application; other groups showed no
reaction to treatment, or a positive reaction. Surface-dwelling microarthropods
appeared to be affected less negatively than were soil-dwelling microarthropods.
Badejo and Van Straalen (1992) tested the effects of atrazine on the growth and
reproduction of the collembolan Orchesella cincta. The lethal concentration (LC50)
for atrazine was estimated at 224 µg/g atrazine in food. Mortality and molting
frequency increased with increasing concentrations of atrazine. The no observed
effect concentration (NOEC) on egg production of O. cincta was 40 µg/g. Based on
data for five collembolan species, 2.7 µg/g was estimated to be the hazardous
concentration for 5% of soil invertebrates, which corresponds to the recommended
field rate of 2.5 µg/g. House et al. (1987) investigated the impact of seven herbicides
on miroarthropods and decomposition. No effect of any herbicide was observed on
numbers of microarthropods, but decomposition of wheat straw was more rapid in
soils without than with herbicide.

Generally, phenoxy acetic acid herbicides (e.g., 2,4-D, 2,4,5-T, 2-methyl-4-
chlorophenoxyacetic acid) do not depress soil fauna directly with toxic effects, but
indirectly through reduced vegetation and smaller additions of organic matter to soil
(Andrén and Lagerlöf, 1983). Simazine, a triazine herbicide, is deleterious to most
soil fauna (Edwards and Stafford, 1979).
Certain compounds such as the fungicide benomyl and its conversion product
carbendazim have negative effects on soil biota even in low concentrations (Andrén
and Lagerlöf, 1983). Applications of the fungicide captan to field soil reduce the
abundance of saprophytic fungi and fungal-feeding mites compared with untreated
field soil (Mueller et al., 1990).
CONCLUDING REMARKS
There are many other factors that influence diversity and function in agricultural
soils. Greater diversity and later successional communities of soil fauna such as
nematodes are found in soils with perennial crops compared with soils with annual
crops (Ferris and Ferris, 1974; Wasilewska, 1979; Freckman and Ettema, 1993;
Neher and Campbell, 1994). Root growth is more extensive and less ephemeral with
perennial than with annual crops. Differences between soils with perennial (e.g.,
meadow fescue Festuca pratensis L.) and annual (e.g., barley) crops may be less
pronounced for perennial crops younger than 3 years old than more mature crops
(Böstrom and Sohlenius, 1986).
In fields where annual crops are grown, the diversity of soil fauna is increased
with management practices such as crop rotation, polycultures, crop mixtures, trap
crops, and intercropping. For example, populations of oribatid (cryptostigmatid) and
prostigmatid mites and springtails were greater in soils with crop rotation than
without (Andrén and Lagerlöf, 1983). However, diversity of nematode communities
in soils in intercropping systems of yellow squash (Cucurbita pepo L.) and cucumber
(Cucumis sativa L.) with alfalfa (Medicago sativa L.) or hairy indigo (Indigofera
hirsuta L.) were not greater consistently than monocultures (Powers et al., 1993).
The lack of consistent difference in diversity was attributed to fluctuations in diver-
sity occurring within the growing season. Further studies are needed to elucidate
the role of faunal diversity in soils with heterogeneous cropping systems.
Agricultural systems are complex, and most research studies have focused on
single factors in an effort to reveal underlying mechanisms. This results in a lack
of understanding of how multiple environmental and biotic factors interact to affect
soil biodiversity and function. As interest in reducing fossil fuel–based inputs
increases, reliance on natural cycles and processes will increase. We should allow
the soil to work for us and not work against it (Elliott and Coleman, 1988). More
research is needed to determine the impact of multiple and interacting management
practices on biodiversity, nutrient cycling, pest populations, and plant productivity.
With this information, we can maximize our ability to tailor agricultural practices
to optimize crop productivity while positively affecting beneficial soil organisms
and the functions they perform.

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CHAPTER 4
Uses of Beneficial Insect Diversity in

Agroecosystem Management
Petr Starý and Keith S. Pike
CONTENTS
Introduction
Beneficial Insects and Their Value
Biodiversity Crisis
Biodiversity in Farmland
Biodiversity Monitoring
Key Beneficials in Agroecosystems
Intraspecific Diversity of Beneficials
Biodiversity of Beneficials in Insect Pest Control Systems
Importation of Beneficials
Natural Enemy Spectrum, Selection, and Adaptation
Beneficial Introductions and Specificity
Conservation of Beneficials
Habitat Management, Crop Structure, and Diversity
Food Sprays and Semiochemicals
Modification of Chemical Pest Control Practices
Augmentation of Beneficials
Releases of Mass-Reared Natural Enemies
Pesticide-Resistant Beneficials
Landscape Ecology
Summary
References

INTRODUCTION
Beneficial Insects and Their Value
Parasitic and predatory insects occur within a wide range of insect groups, and
at times, can be relatively abundant. Some common representatives include predatory
carabid, coccinellid, and staphylinid beetles; predatory bugs; lacewings; syrphid,
chamaemyiid, and other predatory flies; ants; and parasitic wasps. Related to the
beneficial insects are predatory mites and spiders. These different beneficials prey
on and reduce phytophagous pest populations and, thus, promote higher standards
of crop health and economic returns. They can be highly effective at little or no cost,
serving as biotic insecticides in place of chemicals and providing long-term control
without the target pests developing significant resistance to them, and with minimal
or no harm to humans or the environment (Wilson and Huffaker, 1976).
The value or full value of the insect natural enemies is not always realized
because the preferred agent(s) for the target pest(s) are not present, or their abundance
or activity is limited by environmental factors, in particular, by human-implemented
practices such as clean cultivation, pesticide application, etc. (Johnson and Wilson,
1995). Natural enemies do not act in isolation but within the framework of natural
enemy communities comprising individual guilds (Ehler, 1994). These communities
manifest definable structure in which species richness and host range are fundamen-
tal properties (Hawkins and Sheehan, 1994). Individual members often show marked
differences in their utilization of successive life stages of their hosts (Mills, 1994)
and manifest certain positions (trophic levels) in the feeding hierarchy (Powell et
al., 1996). Also, more or less competitive interactions may occur among species
participants (Rosenheim et al., 1995).
The significance of beneficials in agroecosystems is often taken for granted or
overlooked, sometimes when they are most effective. At times their significance
becomes apparent in their absence or when they have been reduced to ineffective
levels allowing the pest to reach crop-injuring levels (Ridgway and Vinson, 1977).
The value is also apparent when exotic pest species in a new area rapidly reach pest
status, but later are suppressed by adapting indigenous natural enemies or newly
released beneficials or both (Clausen, 1978; Nechols et al., 1995).
Biodiversity Crisis
Biodiversity or natural habitat resources are dwindling, in large measure because

of urban and agricultural spread and commercial development (LaSalle and Gauld,
1992). Natural enemies that demonstrate an ability to become community members
in an agroecosystem have, in general, a much better chance to survive compared
with those associated only with natural ecosystems. Through adaptation, at least
some beneficials have overcome or are overcoming the biodiversity crisis by moving
into or between cultivated landscapes. Plant diversity in cultivated landscapes con-
tributes to overall biodiversity, whereas monocultures, especially large-scale mono-
cultures, usually result in fewer species. In all cases, the diversity of beneficials and
their management in agroecosystems should be considered from a dual viewpoint,
inclusive of both the agroecologist and nature conservationist (Samways, 1993).

Biodiversity in Farmland
The elements of biodiversity, both floristic and faunistic, that sustain efficacious
levels of beneficial insects in farmland settings are challenging to obtain, since
farming constitutes a disturbance of the land, and therefore a disturbance of natural
systems and a diminishing of the biotic elements. The greater the disturbance, the
fewer the opportunities for the natural biota to exist. Present trends in biodiversity
development center on equalizing ecological losses through crop diversification,
adjacent landscape preservation, and intentional introduction of biotic agents
(Michal, 1994). Landscape ecology and biodiversity are ecologically connected and
mutually dependent (Carroll, 1990; van Hook, 1994). Overall activities that support
and increase diversity and ecological stability in agroecosystems include, but are
not limited to, development of biocorridors and biocenters, heterogeneous crops and
crop structuring, polycultural crop rotation, biocontrol introductions, and pesticide
use modifications (Paoletti et al., 1992; Petr and Dlouhy, 1992).
Biodiversity Monitoring
Biodiversity is described at three fundamental levels — ecosystem diversity,

species diversity, and genetic diversity (Office of Technology Assessment, 1987).
Changes in the diversity can be monitored by indicator species (Noss, 1990). Mon-
itoring biodiversity in farmlands should account for biota both in the crops and in
the soils. Apart from the common crops and livestock in agriculture, some 200,000
other species of plants and animals are involved in agricultural production and
perform many essential functions, such as nutrient recycling, waste decay, plant
protection, pollination (Pimentel et al., 1989).
Key Beneficials in Agroecosystems
Ideal integrated pest control should reflect ecological approaches that not only
target the pest, but also account for the key natural enemies and associated interac-
tions (LaSalle and Gauld, 1992; LaSalle, 1993). The success or lack of success of
parasitic and predatory species is commonly linked to not only the target host, but
other hosts, bioagents, habitats, and abiotic factors. Understanding the host range,
host preferences, seasonal occurrence, interspecies competition and displacement,
and habitat and food resource requirements of the beneficials is important to safe-
guarding them, increasing their numbers, and enhancing their performance. The
diversity of beneficials in agroecosystems is often linked to natural or undisturbed
environments. Where strong ties exist between biocontrol agents of agriculture and
plant communities of natural diversity, it is important that these are identified and
that the biodiversity linkages are preserved to undergird and support the existence
of the beneficials year-round, and in some cases, for use in redistribution and
introduction elsewhere. Some indigenous beneficials, though less important against
present pests, may be key to preventing future introduced species from becoming
problematic. Environmental diversity should be conserved regardless of what is

known about the taxonomy or biology of the flora and fauna. Beneficials or groups
of beneficials that are active in individual cropping systems commonly move in and
between crops and noncultivated ecosystems. Therefore, any search for beneficials
for introduction purposes should include not only the agroecosystems of interest,
but also the surrounding habitat (Waage, 1991).
Intraspecific Diversity of Beneficials
A common feature of many agroecosystems is a reduction in species richness

coupled with high populations of selected other species. With parasitic Hymenoptera,
for example, the phenomenon may affect not only the interspecific relationships
within certain parasitic spectra, but also the intraspecific diversity of individual
parasitoids (Unruh and Messing, 1993). There may be changes in the gene flow
between populations from different host species as well as certain dominance of
features in populations from highly populated dominant hosts. These factors are
important in the drift of parasitoids between different hosts. One of the main prob-
lems in agroecosystems is the lack of hosts for host alternation and relationships
(Nemec and Starý, 1984; 1985; Powell, 1986).
Biodiversity of Beneficials in Insect Pest Control Systems
Biodiversity as a factor in pest control varies widely between countries and areas
of the world. Introduction strategies in classical biological control typically center
on the full range of beneficials attacking the target pest throughout the world, with
the aim to find, select, and use the most promissive agents from the world complex.
The introduction of broad-spectrum pesticides starting with DDT contributed to
a strong demand for pest-free crops. Resistance to pesticides and pest resurgence
connected with natural enemy losses, however, led to the development of integrated
pest management (IPM) (Stern et al., 1959; van den Bosch and Stern, 1962; Smith
and Reynolds, 1966) and, more recently, alternative pest management emphasizing
ecologically adapted and biorationally based approaches to the exclusion of synthetic
pesticides uses (U.S. National Research Council, 1989; Vereijken, 1989).
Additionally, sustainable agriculture initiatives stimulated efforts to increase and
maintain greater biodiversity through landscape protection of fauna and flora, e.g.,
introduction of grassland meadows in place of arable land (Petr and Dlouhy, 1992).
Diversity, its support and enhancement through species richness, rotations, inter-
cropping, cover crops, etc., is one of the basic principles of agroecology in sustain-
able agriculture systems (Thrupp, 1996).
IMPORTATION OF BENEFICIALS
Natural Enemy Spectrum, Selection, and Adaptation
The introduction of exotic natural enemies to control exotic pests is the primary
approach in classical biological control (DeBach, 1964). Exotic pests, usually

inadvertently introduced and without their natural enemies, commonly result in rapid
increases and population outbreaks. The lack of natural enemies provides the impetus
for purposeful introductions of potentially promising natural enemies. Such enemies
are usually sought in the homeland of the target pest, but sometimes are obtained
from areas outside the original home of the pest, where secondary adaptation by
indigenous species has occurred. Consideration of world populations broadens the
scope for selection of useful species and biotypes, and increases the prospect for
success (Starý, 1970a; Huffaker et al., 1971; Greathead, 1986; Ehler, 1990; 1992).
New beneficial introductions add to the biodiversity of pest-infested areas, but
also they can impact the indigenous biota and, in some instances, stand as biotic
contaminants (Samways, 1993) in terms of host displacement and nontarget prey
interspecies competition. Ideally, environmental impact determinations or projections
on prey specificity should be studied prior to a release (Starý, 1993). With the excep-
tion of strict monophagous agents, some adaptation to indigenous biota is expected.
Endemic ecosystems may be disrupted or partially disrupted due to various reasons
and thus are more easily attacked by exotic invaders (Starý, 1994). An introduced
biocontrol agent may not be necessarily harmful to indigenous communities where
the spectrum of natural enemies includes numerous broadly oligophagous species
(Starý et al., 1988) or where invasive species are suppressed (Samways, 1993).
New pests in an area can be attacked and suppressed to varying degrees by
indigenous natural enemies. Here, there is no increase in species richness of natural
enemies, simply expanded adaptation and enlargement of the prey/host spectrum.
However, where new strains or biotypes occur, differences in intraspecific compe-
tition may result.
Once introduced species become established, they become part of the natural
enemy spectrum for the target pest and associated area, and thus subject to all of
the management approaches applied, such as conservation, augmentation, etc. New
agents, however, need to be classified as generally more vulnerable to environmental
modifications depending upon the degree of adaptation in their new environment.
Beneficial Introductions and Specificity
Classical biological control efforts are seldom instituted until a pest outbreak
occurs, and, even then, there can be a delay because of the time involved to search
for, import, quarantine, mass-rear, and release new agents. Although not generally
followed, another approach is preventative biocontrol, the introduction of promising
exotic biocontrol agents prior to the appearance of a forecasted or expected target
pest. Except for strict monophages, oligophagous agents may be introduced for
establishment on alternate prey/host species in association with the target agroeco-
system (Starý et al., 1993). These then are present to attack the arriving pest,
possibly before it is detected by humans. The approach provides a temporal advan-
tage, and more or less limits population outbreak of the invading pest. Switching
from a native host to a related introduced species can occur with striking results,
and may include the indigenous beneficials (LaSalle, 1993). Understanding the host
range of introduced exotics is key to achieving success in preventative biocontrol
(Starý et al., 1993).

The host range of introduced beneficials should always be considered, i.e.,
control of more than one pest using the same regulatory agent. This requires an
oligophagous agent. In principle, oligophagous agents can attack several pests in
the same or different crops (Starý et al., 1993). Such situations contribute to the
stability of the natural enemy interactions across cropping systems. However, such
introductions are not without risks, e.g., host preference, poor alternation, or species-
specific strains may eventually develop. The strategies, necessary attributes, and
opinions for introducing biocontrol agents are widely discussed in the literature
(DeBach, 1964; Starý, 1970a; Huffaker et al., 1971; van den Bosch and Messenger,
1973; Clausen, 1978; Croft, 1990; Ehler, 1990; Miller, 1993; Nechols et al., 1995).
CONSERVATION OF BENEFICIALS
Conservation in this discussion means modifying any environmental factors that

are adverse to beneficials (DeBach, 1964) and adding requisites (McMurtry et al.,
1995). This is a type of environmental insect control (Stern, 1981), a manipulation
of the ecosystem to make it less favorable to the pest and more favorable to the
natural enemies resulting in reduced pest levels (Mayse, 1983). Such approaches
need to be considered in the context of the whole environment, the agroecosystem
(target and adjoining crops) and surroundings, as these often have mutual connec-
tions. Many insects, whether classified as pests, beneficials, or indifferents, exhibit
population drift (Starý, 1978; Bosch, 1987; Vorley and Wratten, 1987; van Emden,
1988). The boundary zone or ecotone where individual crops and noncrops overlap
is frequently essential to the conservation and management of beneficials, both
indigenous and introduced (DeBach, 1964; van Emden, 1965; Ridgway and Vinson,
1977; Stern, 1981; Powell, 1986; Gross, 1987; Martis, 1988; Altieri et al., 1993;
Samways, 1993; Johnson and Wilson, 1995; McMurtry et al., 1995). Strict separation
of ecosystems does not occur in nature.
Habitat Management, Crop Structure, and Diversity
Habitat management is viewed as a strategy aimed at designing and constructing

“phytocenotic architecture” dominated by plants that support populations of natural
enemies (Altieri and Whitcomb, 1979; Altieri, 1983). Diversification of habitat is
achieved through crop structure, protective refugia, occurrence of alternative
prey/host, and supplementary food resources (nectar, pollen). Crop structure is the
agroecosystem and its specific characteristics, its biotic composition, seasonality,
etc. Protective refugia are defined as habitats in which beneficials can survive critical
periods of the year (principally summer and winter periods) to disperse later to
crops. Protected refugia can include a wide array of plant types and setting, e.g.,
rangeland, weedy field margins, autumn-sown crops, etc.
Alternative prey/hosts and their availability or proximity to crops are important
at times when the target pest is low in numbers. Alternative hosts can improve the
synchrony between natural enemies and the target pests. For a given beneficial, the
alternative host might be a nonpest species feeding on wild plants or it might be a

pest species different from the target on another crop (van den Bosch and Telford,
1964; Powell, 1986). For some aphid parasitoids, utilization of alternative hosts in
proximity to the target pest and crop is known as multilateral control or the multi-
lateral control approach (Starý, 1972; 1978). Such an approach takes advantage of
the oligophagous host range of the bioagent. Switching from one host to another
can be effective or ineffective depending upon the biotype or species specificity of
the bioagent (Gordh, 1977). Population genetics in relation to host alternation is
currently of high research interest. Until recently, host alternation was based solely
on field observation and laboratory transfers; now, studies on population molecular
genetics are further clarifying the species or species strains of key importance and
their host alternation dynamics (Unruh et al., 1983; Nemec and Starý, 1985).
Sources of food such as nectar and pollen are requisite for hymenopterous
parasitoid (van Emden, 1962) and syrphid (Hickman and Wratten, 1996) adults to
ensure effective reproduction. More often than not, flowering plants in or even around
agriculture crops for such uses are not always readily available (van Emden, 1962;
Altieri and Whitcomb, 1979; Altieri and Letourneau, 1982; Powell, 1986).
The composition, seasonality, field size, and location of crops and noncrops all
affect biodiversity. Semiperennial and perennial crops are generally classified as
more stabilizing for biotic diversity in comparison with annuals; nonetheless, pop-
ulation drift can take place across all settings (Gross, 1987; Andow, 1991). Gliessman
(1987) reported that whenever two or more crops are planted together, there is
increased potential for species interactions, and this would include beneficials.
Diverse landscape mosaics in and around small-sized fields enhance the chances
for greater diversity of beneficials. Nonetheless, relatively high biodiversity is
thought possible even in intensely cultivated areas as long as crops are arranged
together with patches of natural or seminatural areas (Duelli et al., 1989; 1990).
Among the most difficult environments for biocontrol to succeed in are the annual
crop monocultures. These usually lack the resources for the natural enemies to be
efficacious, are grown using cultural practices (e.g., mowing of alfalfa) that often
damage the natural enemy populations, and are present for only part of the year
(Rabb et al., 1976; Powell, 1986). In such circumstances, it may be necessary through
habitat management to maintain small populations of the target pest to ensure
survival of the key beneficials (Powell, 1986). Volunteer crops along field margins
may be useful in this regard.
Preservation, establishment, or sustainment of small heterogeneous strip habitats,
in or neighboring crop farmlands, adds to the overall biodiversity of farm or area
(Nentwig, 1993). Also, strip farming, especially intercropping, increases natural
enemy opportunities for predation and parasitism over that of strict monocultures
(Powell, 1986).
Cultural practices such as full field mowing of perennial legumes and grassy
meadows hinder bioagent survival and success. These negative effects can be par-
tially offset by strip mowing which affords arthropods an opening to retreat to uncut
portions (Stern et al., 1964; Müller-Ferch and Mouci, 1995) and, thus, increases or
strengthens the population stability of the beneficials (Schlinger and Dietrick, 1960;
van den Bosch et al., 1967; Starý, 1970b).

Heterogeneous herbaceous strips constitute more suitable habitat for field species
than strips of shrubs or trees that tend to harbor ecologically different forest species
(Nentwig, 1988). Selected weed species used in strips within crop fields have been
shown to attract and aid in the conservation of beneficials (Starý, 1964; van Emden,
1965; Gliessmann, 1987; Nentwig, 1988; 1992; 1993; 1994; 1995; Frei and Manhart,
1992; Weiss and Nentwig, 1992; Hausmann, 1996). Wyss (1995) showed that in
apple orchards certain flowering weed strips resulted in more aphidophagous pred-
ators and fewer aphid pests than like areas without weeds. Nectar-bearing plants
(Chumakova, 1977) and rich undergrowth of wild flowers (Leius, 1967) showed
similar beneficial effects. Many natural enemies occur commonly in association with
wild or natural habitat not always classified as weeds (van Emden, 1965). Tillage
practices (no-till, minimum, conventional), tillage timing, mowing, or other agro-
nomic practices can influence, sometimes significantly, the performance, success,
maturation, and dispersal of beneficials (Bugg and Ellis, 1990; Bugg, 1992). Molthan
and Ruppert (1988) demonstrated that flowers in wide boundary strips attracted
beneficials, some being especially attractive and nutritionally suitable. They further
recommended protection or arrangement of boundary strips in the framework of
agricultural extensification.
Cultures of medicinal, culinary, and ornamental herbs, such as sweet fennel
(Foeniculum vulgare) and spearmint (Menta spicata), grown in organic market
gardens near various vegetable and tree crops are known to attract several adult
entomophagous Hymenoptera and flower-visiting beneficials (Sawoniewicz, 1973;
Bugg and Wilson, 1989; Bugg et al., 1989; Maingay et al., 1991; Bugg and Wad-
dington, 1994). Some studies on herb attractiveness have listed not only the sampled
species, but also have determined or attempted to determine their significance and
effect on pests in nearby crops (Bugg and Waddington, 1994 ).
Pollen can serve as a supplemental or essential food source for beneficials. For
example, Ouyang et al. (1992) report that pollen can positively affect polyphagous
predacious mites, especially during periods when their arthropod prey is scarce.
Green manure crops such as faba bean (Vicia faba) can manifest similar positive
effects on beneficials (Bugg and Ellis, 1988; Bugg et al., 1989). And, habitat manip-
ulations, such as the addition of mulch and flowers, may enhance spider densities
and lower the number of pest insects in a mixed vegetable system (Riechert and
Bishop, 1990).
Cover crops in general are known to affect a number of phenomena in orchards.
They may harbor pest species, but they can also lead to increased numbers of insect
natural enemies and heightened pest biocontrol (Bugg et al., 1990; Bugg and Wad-
dington, 1994). Some covers, or mixtures of cover crops, constitute field insectaries
and may be marketable as “insectary crops” or crops that support high densities of
beneficials. Different covers may require different management protocols, depending
upon whether they are supplementary (alternative prey or hosts, pollen) or comple-
mentary (nectar, honeydew) food sources (Bugg, 1992). For a detailed review of
cover crop management in temperate zone orchard crops (almond, pecan, walnut,
apple, cherry, peach, and citrus), see Bugg and Waddington (1994).

Field margins and crop edges can be highly supportive of beneficials, but their
relative abundance varies depending upon the mix of plants at the margins and the
adjacent crop (Dennis and Fry, 1992). Some studies have shown that field margins
can increase the diversity of arthropods within the crop and that movement between
the margin and the field can be significant. Margin habitats provide the stability
needed for species that would otherwise not survive across all seasons. The landscape
matrix of field margins can be vital for effective field dispersal and conservation
success (Röser, 1988; Dennis and Fry, 1992).
Crop edges adjacent to hedges and broad grassy strips bear a rich fauna of
beneficials and should not be treated with pesticides or receive fertilizer (van den
Bosch and Messenger, 1973; Morris and Weeb, 1987; Basedow, 1988; Holtz, 1988;
Klingauf, 1988; Welling and Kokta, 1988). Dover (1991) and Samways (1993)
showed that 6-m-wide edges around cereal fields receiving reduced and selective
pesticides helped conserve beneficials. Welling and Kokta (1988) reported that wide
headlands with a large source of flowering plants guaranteed nutrition for flower-
visiting beneficials, served as a refuge for different species (before and after harvest),
and acted as a bridge between isolated biotypes.
The plants that compose hedges, hedgerows, and windbreaks vary widely, as does
their significance as reservoirs for beneficials (Solomon, 1981). Wide hedgerows or
windbreaks composed of trees and shrubs appear to function as a type of biocorridor
across the landscape (Forman and Baudry, 1984); the associated beneficials are in
part forest-edge species. The hedgerows and windbreaks, together with boundary
strips and unsprayed field margins, represent a functional part of the agroecosystem,
influencing positively the beneficials (Knauer, 1988), and thus should be encouraged
on farms (Basedow, 1988). A number of papers cover hedgerow/windbreak habitats
in detail, including the role of beneficials and IPM (Lewis, 1969; Zwölfer et al., 1984;
Stechman and Zwölfer, 1988; Welling et al., 1988; Häni, 1989).
Food Sprays and Semiochemicals
Food requirements of predaceous species sometimes vary between life stages.

Larval stages may be carnivores, while adults may feed on nectar, honeydew, and
pollen. For hymenopterous parasitoids, nectar and pollen requirements are common
for adults, but not for parasitic larvae. With some crops, nectar, pollen, and honeydew
sources are insufficient or unavailable. By providing an artificial food supplement,
beneficials may be retained, arrested, or stimulated to oviposit. Treatments may
consist of yeast, sucrose solution, or artificial honeydew (Hagen and Bishop, 1979;
Gross, 1987).
Natural enemies are known to respond to a number of environmental cues in the
course of locating desired habitats, plants, prey/hosts, and the opposite sex. Behavior-
controlling chemicals (semiochemicals) are rather species specific. In theory, syn-
thetically derived semiochemicals may be used to attract increased numbers of
natural enemies into a crop, prolong their searching activity, and improve their
performance (Vinson, 1977; 1981; Lewis, 1981; Nordlund et al., 1981; Powell, 1986;
Gross, 1987; McMurtry et al., 1995).

Modification of Chemical Pest Control Practices
Biodiversity of natural enemies in agroecosystems may be substantially affected

by the use of pesticides. Nonselective treatments are toxic to beneficials. They
decrease populations, contributing to pest outbreak. For this reason and others,
pesticides have begun to be used with greater care. Emphasis is beginning to center
more on the development and use of selective pesticides, on target-directed appli-
cations, and on applications timed to avoid the direct treatment of beneficials.
Information on pesticidal effects on beneficials is extensive.
In some respects, the value of beneficials has been heightened by the overuse
of pesticides leading to chemical resistance, secondary pest outbreaks, and environ-
mental pollution. IPM concepts and strategies, from the earliest discussions, have
centered on selective treatments to protect beneficials as key components in inte-
grated control (Stern et al., 1959; Smith and Reynolds, 1966; Croft, 1990; McMurtry
et al., 1995).
Nonselective pesticides directly and negatively affect natural enemies, sometimes
even the larval stages within a host, e.g., parasitoids. Pesticides, both selective and
nonselective, indirectly impact beneficials by diminishing prey/host populations and,
in turn, force surviving beneficials to disperse to other communities to find food
(Petr and Dlouhy, 1992). In a few cases, pesticide-affected systems have led to
changes in gene diversity of beneficials through selection of pesticide-resistant
strains.
Biocontrol is not the primary approach for some agroecosystems, and may never
be, but it is or could be the key component in many agroecosystems. In most natural
environments, biocontrol provides common, if not perennial regulation. With the
removal of pesticides, diversity of beneficials and restoration of biocontrols are
possible (Hagen et al., 1971). Where market demands require blemish-free products
or where farm economics dictate treatment to protect investments, chemicals will
likely remain a standard defense.
Conservation activities targeting beneficials through habitat diversification may
be adversely disrupted by herbicides. Way and Cammell (1981) suggested that insect
communities, including natural enemies, in and around agroecosystems are affected
more by herbicides than by pesticides. Similarly, fertilizer, especially in heavy
dosage, can adversely affect conservation efforts.
AUGMENTATION OF BENEFICIALS
Augmentation, broadly defined, covers all of the activities that improve the
effectiveness of beneficials (DeBach, 1964), such as new species releases (inocula-
tive or inundative), planned genetic change, landscape modification, and so on. The
addition of a new species or strain of species into a new area increases species
diversity, but it can also affect interspecific relationships and population genetic
characteristics.

Releases of Mass-Reared Natural Enemies
There are two main types of releases, inoculative and inundative. Inoculative
releases utilize species intended for permanent establishment, where progeny of each
successive generation continue as regulatory agents. Inundative releases utilize spe-
cies intended for short-term action (permanent establishment not expected) and act
as biotic insecticides.
Pesticide-Resistant Beneficials
Discovery, mass-rearing, and redistribution of endemic resistant populations of

beneficials against target pests, or laboratory-planned genetic alteration, hybridiza-
tion, or artificial selection for pesticide-resistant beneficials and subsequent field
release, are measures intended to amplify existing levels of biocontrol. The use of
such agents is expanding rapidly. Common cases include predatory mites, aphid
predators, and parasitoids in deciduous tree fruit and nut crops (Hoy, 1985; 1995;
Croft, 1990; Johnson and Wilson, 1995; McMurtry et al., 1995).
Landscape Ecology
Land management planning and IPM efforts must consider biocontrol issues,
the players, their role, protection, and sustainment. Agroecosystems are linked not
only to the surrounding natural lands, but also to biological diversification from
distant sources (van Hook, 1994). Beneficial insect diversity needs to be dealt with
as part of the overall diversity of the landscape. Biodiversity exists in a matrix of
habitat patches including managed and natural environments. The landscape and
beneficials are ecologically linked and interdependent (Carroll, 1990; van Hook,
1994).
IPM programs, though strongly based on ecological principles, still rely heavily
on reductionistic approaches to control single pests on single crops (van Hook, 1994).
Discussions on beneficials often center on the interdependency of specific habitats.
Although it may not be possible to view all of the biological peculiarities of an
ecosystem or to make overall generalizations, greater accounting of multiple factor
linkages and features in common (e.g., epigean fauna, role of flowering plants,
stability of plants, crop culture specifics, etc.) are necessary to appropriately under-
stand, conserve, and manage beneficials in relation to their phytophagous hosts.
Some beneficials may be linked to their prey/host for only a part of their life cycle.
Altieri (1983) divided the crop landscape into two groups: (1) heterogeneous
(crop, meadow, woodland, windbreak, etc.) and (2) homogeneous (continuous crop
cultures). The latter could be further subdivided into extensive plantings of annual
monocultures, (e.g., small grains), semiperennials (e.g., alfalfa), perennials (e.g.,
grapes), or mixtures of these in small to moderate size.
The most effective beneficials in agriculture are usually species that are peren-
nially present in farmlands. Long-term agricultural landscape optimization should

support equilibrium between pest and beneficial, achievable in part or in total, through
extended diversification of crops and wild plants, or through development of biocor-
ridors adjoining particular managed habitats (Petr and Dlouhy, 1992). Farmland
optimization will vary broadly between areas, but certainly will require long-term
ecological planning and commitment with the flexibility to accommodate new dis-
coveries for improved activity and perennial sustainment of desired beneficials.
Today, high-yielding, high-quality, insect-, disease-, and weed-free monocultures
require high-energy inputs to produce and harvest. With production costs escalating,
interest in enduring and efficient alternatives with less dependence on fossil fuels is
escalating (Gross, 1987). Central to this is the management of agricultural commu-
nities (Altieri, 1981; Altieri et al., 1993), where selective diversity in, or in association
with, cropping systems is used to enhance the stability, survival, and efficiency of
the natural community inhabitants (Gross, 1987).
SUMMARY
Natural community beneficials in cropping systems are interlinked with sur-

rounding vegetation matrices. Their biodiversity is affected by numerous farmland
approaches and activities. But much of the biological aspects of the beneficials, their
specific interrelationships, differences in relative abundance, performance, and man-
agement in and between agroecosystems, geographical areas, climatic zones, etc.
are yet to be determined or determined in detail. The ecological planning, selection,
adaptation, and management of beneficials in a given crop may initially appear costly
if just the direct immediate system inputs and profits are considered (Dahlberg,
1992). The measures to enhance diversity and efficiency of beneficials can be
difficult, and may not always be feasible. For this reason, each agricultural situation
must be assessed separately (Altieri et al., 1993).
Management of beneficials based on current knowledge must be considered as
developmental for now. Changes in ecological optimization and stability concepts
and implementation will follow further crop structure analysis and landscape diver-
sification research, on-farm trials, and successful IPM examples under variable
environments. Legume-based crop rotations, recognized as stabilizing crops, used
to improve soils and crop yields, may serve to increase the diversity of beneficials,
and may play a significant role in the bilateral drift of beneficials to annuals.
The effects of natural enemies on target pests have economic implications. It is
possible to evaluate numerically the ecological value of certain species within a
complex net of relationships and connections. We may not appreciate or understand
the role or the importance of a given species until after its disappearance, and there
may be some insects or insect groups not directly determined to be beneficials in
an economic sense, but that may contribute in other ways, such as to the beauty of
the environment (Martis, 1988). This could occur in natural systems and in agroec-
osystems via landscape diversification. Butterflies flying over flowering meadows,
hedgerows, weedy ecotones, roadsides, or similar biocorridors improve the often

tiring effect of visually plain, monotonous monocultures. Simply stated, biodiversi-
fication can be used to improve the aesthetics of the farmland.
This chapter is not an exhaustive review. We have highlighted some key factors,
outlined some of the complexities involved in the biology, pathways, and population
dynamics of insect natural enemies and their management, cited references for more
detail, and mentioned in brief the need for further research.
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schaftspflege, 3(3).

CHAPTER 5
Biodiversity, Ecosystem Function, and

Insect Pest Management in
Agricultural Systems
Miguel A. Altieri and Clara I. Nicholls
CONTENTS
Introduction
The Nature and Function of Biodiversity in Agroecosystems
Patterns of Insect Biodiversity in Agroecosystems
Plant Biodiversity and Insect Stability in Agroecosystems
Patterns of Landscape Structure and Insect Biodiversity
Conclusion
References
INTRODUCTION
Today, scientists worldwide are increasingly starting to recognize the role and
significance of biodiversity in the functioning of agricultural systems (Swift et al.,
1996). Research suggests that, whereas in natural ecosystems the internal regulation
of function is substantially a product of plant biodiversity through flows of energy
and nutrients and through biological synergisms, this form of control is progressively
lost under agricultural intensification and simplification, so that monocultures in
order to function must be predominantly subsidized by chemical inputs (Swift and
Anderson, 1993). Commercial seedbed preparation and mechanized planting replace
natural methods of seed dispersal; chemical pesticides replace natural controls on

populations of weeds, insects, and pathogens; and genetic manipulation replaces
natural processes of plant evolution and selection. Even decomposition is altered
since plant growth is harvested and soil fertility maintained, not through nutrient
recycling, but with fertilizers.
One of the most important reasons for maintaining and/or encouraging natural
biodiversity is that it performs a variety of ecological services (Altieri, 1991). In
natural ecosystems, the vegetative cover of a forest or grassland prevents soil erosion,
replenishes groundwater, and controls flooding by enhancing infiltration and reduc-
ing water runoff. In agricultural systems, biodiversity performs ecosystem services
beyond production of food, fiber, fuel, and income. Examples include recycling of
nutrients, control of local microclimate, regulation of local hydrological processes,
regulation of the abundance of undesirable organisms, and detoxification of noxious
chemicals. These renewal processes and ecosystem services are largely biological;
therefore, their persistence depends upon maintenance of biological diversity. When
these natural services are lost as a result of biological simplification, the economic
and environmental costs can be quite significant. Economically, in agriculture the
burdens include the need to supply crops with costly external inputs, since agroec-
osystems deprived of basic regulating functional components lack the capacity to
sponsor their own soil fertility and pest regulation. Often the costs involve a reduction
in the quality of the food produced and of rural life in general due to decreased soil,
water, and food quality when erosion and pesticide and/or nitrate contamination
occurs (Altieri, 1995).
Nowhere are the consequences of biodiversity reduction more evident than in
the realm of agricultural pest management. The instability of agroecosystems
becomes manifest as the worsening of most insect pest problems is increasingly
linked to the expansion of crop monocultures at the expense of the natural vegetation,
thereby decreasing local habitat diversity (Altieri and Letourneau, 1982; Flint and
Roberts, 1988). Plant communities that are modified to meet the special needs of
humans become subject to heavy pest damage, and generally the more intensely
such communities are modified, the more abundant and serious the pests. The effects
of the reduction of plant diversity on outbreaks of herbivore pests and microbial
pathogens are well documented in the agricultural literature (Andow, 1991; Altieri,
1994). Such drastic reductions in plant biodiversity and the resulting epidemic effects
can adversely affect ecosystem function with further consequences on agricultural
productivity and sustainability (Figure 1).
In modern agroecosystems, the experimental evidence suggests that biodiversity
can be used for improved pest management (Altieri and Letourneau, 1984; Andow,
1991). Several studies have shown that it is possible to stabilize the insect commu-
nities of agroecosystems by designing and constructing vegetational architectures
that support populations of natural enemies or that have direct deterrent effects on
pest herbivores (Perrin, 1980; Risch et al., 1983). This chapter analyzes the various
options of agroecosystem design which, based on current agroecological theory,
should provide for the optimal use and enhancement of functional biodiversity in
crop fields.

Figure 1 The influence of intensification on biodiversity and function in agricultural ecosystems
as it relates to the role of arthropod biodiversity. (Modified from Swift and Anderson,
1993.)
THE NATURE AND FUNCTION OF BIODIVERSITY IN

AGROECOSYSTEMS
Biodiversity refers to all species of plants, animals, and microorganisms existing

and interacting within an ecosystem. In agroecosystems, pollinators, natural enemies,
earthworms, and soil microorganisms are all key biodiversity components that play
important ecological roles, thus mediating such processes as genetic introgression,
natural control, nutrient cycling, decomposition, etc. (Figure 2). The type and abun-
dance of biodiversity in agriculture will differ across agroecosystems which differ
in age, diversity, structure, and management. In fact, there is great variability in
basic ecological and agronomic patterns among the various dominant agroecosys-
tems. In general, the degree of biodiversity in agroecosystems depends on four main
characteristics of the agroecosystems (Southwood and Way, 1970):
1. The diversity of vegetation within and around the agroecosystem;

2. The permanence of the various crops within the agroecosystem;
3. The intensity of management;
4. The extent of the isolation of the agroecosystem from natural vegetation.
In general, agroecosystems that are more diverse, more permanent, isolated, and
managed with low input technology (i.e., agroforestry systems, traditional polycul-
tures) take fuller advantage of work done by ecological processes associated with

Figure 2 The components, functions, and enhancement strategies of biodiversity in agroecosystems. (From Altieri, M. A., Biodiversity and Pest Management
in Agroecosystems, Haworth Press, New York, 1994. With permission.)

higher biodiversity than do highly simplified, input-driven, and disturbed systems
(i.e., modern row crops and vegetable monocultures and fruit orchards) (Altieri,
1995).
All agroecosystems are dynamic and subject to different levels of management
so that the crop arrangements in time and space are continually changing in the face
of biological, cultural, socioeconomic, and environmental factors. Such landscape
variations determine the degree of spatial and temporal heterogeneity characteristic
of agricultural regions, which in turn conditions the type of biodiversity present, in
ways that may or may not benefit the pest protection of particular agroecosystems.
Thus, one of the main challenges facing agroecologists today is identifying the types
of biodiversity assemblages (either at the field or landscape level) that will yield
desirable agricultural results (i.e., pest regulation). This challenge can only be met
by further analyzing the relationship between vegetation diversification and the
population dynamics of herbivore and natural enemy species, in light of the unique
environment and entomofauna of each and the diversity and complexity of local
agricultural systems.
According to Vandermeer and Perfecto (1995), two distinct components of bio-
diversity can be recognized in agroecosystems. The first component, planned bio-
diversity, is the biodiversity associated with the crops and livestock purposely
included in the agroecosystem by the farmer, and which will vary depending on
management inputs and crop spatial/temporal arrangements. The second component,
associated biodiversity, includes all soil flora and fauna, herbivores, carnivores,
decomposers, etc. that colonize the agroecosystem from surrounding environments
and that will thrive in the agroecosystem depending on its management and structure.
The relationship of both biodiversity components is illustrated in Figure 3. Planned
biodiversity has a direct function, as illustrated by the bold arrow connecting the
planned biodiversity box with the ecosystem function box. Associated biodiversity
also has a function, but it is mediated through planned biodiversity. Thus, planned
biodiversity also has an indirect function, illustrated by the dotted arrow in the figure,
which is realized through its influence on the associated biodiversity. For example,
the trees in an agroforestry system create shade, which makes it possible to grow
only sun-tolerant crops. So the direct function of this second species (the trees) is
to create shade. Yet along with the trees might come small wasps that seek out the
nectar in the tree flowers. These wasps may in turn be the natural parasitoids of
pests that normally attack the crops. The wasps are part of the associated biodiversity.
The trees, then, create shade (direct function) and attract wasps (indirect function)
(Vandermeer and Perfecto, 1995).
The key is to identify the type of biodiversity that is desirable to maintain and/or
enhance in order to carry out ecological services, and then to determine the best
practices that will encourage the desired biodiversity components. As shown in
Figure 4, there are many agricultural practices that have the potential to enhance
functional biodiversity, and others that negatively affect it. The idea is to apply the
best management practices in order to enhance and/or regenerate the kind of biodi-
versity that can subsidize the sustainability of agroecosystems by providing ecolog-
ical services such as biological pest control, nutrient cycling, water and soil conser-
vation, etc.

Figure 3 The relationship between planned biodiversity (that which the farmer determines,
based on management of the agroecosystems) and associated biodiversity and how
the two promote ecosystem function. (Modified from Vandermeer and Perfecto,
1995.)
PATTERNS OF INSECT BIODIVERSITY IN AGROECOSYSTEMS
Arthropod diversity has been correlated with aspects of plant diversity in agro-
ecosystems. A greater variety of plants conforming to a particular crop pattern should
lead to a greater variety of herbivorous insect species, and this in turn should
determine a greater diversity of predators and parasites (Figure 5). A greater total
biodiversity can then play a key role in optimizing agroecosystem processes and
function (Altieri, 1984).
Several hypotheses can be offered to support the idea that diversified cropping
systems encourage higher arthropod biodiversity (Altieri and Letourneau, 1982):
1. Heterogeneity hypothesis. Complex crop habitats support more species than simple
crop habitats; architecturally more complex species of plants and heterogeneous
plant associations, with greater biomass, food resources, variety and temporal
persistence, have more associated species of insects than do architecturally simple
crop plants or crop monocultures on an area-for-area basis. Apparently both species
diversity and plant structural diversity are important in determining insect species
diversity.
2. Predation hypothesis. The increased abundance of predators and parasites in rich
plant associations (Root, 1973) reduce prey densities, at times to such low levels
that competition among herbivores should be reduced. This reduced competition
should allow the addition of more prey species, which in turn support new natural
enemies.
3. Productivity hypothesis. Research has shown that in some situations crop polycul-
tures yield more than monocultures (Francis, 1986; Vandermeer, 1989). This greater

Figure 4 The effects of agroecosystem management and associated cultural practices on the
biodiversity of natural enemies and the abundance of insect pests.
productivity can result in greater insect diversity as the number of food resources
available for herbivores and natural enemies increases.
4. Stability and temporal resource-partitioning hypothesis. This hypothesis assumes
that primary production is more stable and predictable in polycultures than in
monocultures. This stability of production, coupled with the spatial heterogeneity
of complex crop fields, should allow insect species to partition the environment
temporally as well as spatially, thereby permitting the coexistence of more insect
species.
Further research is needed to clarify whether insect species diversity parallels

diversity of vegetation and the productivity of the plant community or simply reflects
the spatial heterogeneity arising from the mixing of plants of different structures.
There are several environmental factors that influence the diversity, abundance,
and activity of parasitoids and predators in agroecosystems: microclimatic condi-
tions, availability of food (water, hosts, prey, pollen, and nectar), habitat requirements
(refuges, nesting and reproduction sites, etc.), intra- and interspecific competition
and other organisms (hyperparasites, predators, humans, etc.). The effect of each of
these factors will vary according to the spatial and temporary arrangement of crops

Figure 5 The relationship between plant and arthropod biodiversity and agroecosystem pro-
cesses. Arrow widths indicate the relative amount of information available on each
link; for example, more work has been done on the responses of herbivore popula-
tions to plant diversity than on the converse.
and the intensity of crop management, as these features affect the environmental
heterogeneity of agroecosystems in several ways (van den Bosch and Telford, 1964).
Although natural enemies seem to vary widely in their response to crop distribu-
tion, density, and dispersion, experimental evidence suggests that structural (i.e., crop
diversity, input levels, etc.) attributes of agroecosystems influence parasitoid and
predator diversity and dynamics. Several of these attributes are related to biodiversity
and most are amenable to management (i.e., crop sequences and associations, weed
diversity, genetic diversity, etc.). Based on the available information, natural enemy
biodiversity can be enhanced and effectiveness improved in the following ways (van
den Bosch and Telford, 1964; Rabb et al., 1976; Altieri and Whitcomb, 1979):
• Multiple introductions of parasitoids and predators through augmentative releases

for biological control;
• Reducing direct mortality by eliminating pesticide use;
• Provision of supplementary resources other than hosts/prey;
• Increasing adjacent and within-field vegetational diversity;
• Manipulating architectural, genetic, and chemical attributes of host plants;
• Use of semiochemicals (behavioral chemicals such as kairomones) to stimulate
host/prey searching behavior and natural enemy retention in the field.
PLANT BIODIVERSITY AND INSECT STABILITY

IN AGROECOSYSTEMS
From the early 1970s on, the literature provides hundreds of examples of exper-
iments documenting that diversification of cropping systems often leads to reduced

herbivore populations (Andow, 1991; Altieri, 1994). Most experiments that have
mixed other plant species with the primary host of a specialized herbivore show
that, in comparison with diverse crop communities, simple crop communities have
greater population densities of specialist herbivores (Root, 1973; Cromartie, 1981;
Risch et al., 1983). In these systems, herbivores exhibit greater colonization rates,
greater reproduction, higher tenure time, less disruption of host finding, and lower
mortality by natural enemies.
There are various factors in crop mixtures that help constrain pest attack. A host
plant may be protected from insect pests by the physical presence of other plants
that may provide a camouflage or a physical barrier. Mixtures of cabbage and tomato
reduce colonization by the diamondback moth, while mixtures of maize, beans, and
squash have the same effect on chrysomelid beetles. The odors of some plants can
also disrupt the searching behavior of pests. Grass borders repel leafhoppers from
beans, and the chemical stimuli from onions prevent carrot fly from finding carrots
(Altieri, 1994).
Alternatively, one crop in the mixture may act as a trap or decoy — the “flypaper
effect.” Strips of alfalfa interspersed in cotton fields in California attract and trap
Lygus bugs. There is a loss of alfalfa yield, but this represents less than the cost of
alternative control methods for the cotton. Similarly, crucifers interplanted with
beans, grass, clover, or spinach are damaged less by cabbage maggot and cabbage
aphid. There is less egg laying on the crucifers, and the insect pests are subject to
increased predation (Altieri, 1994).
The two hypotheses that have been proposed to explain lower herbivore abun-
dance in polycultures, the resource concentration hypothesis and the enemies hypo-
thesis (Root, 1973), identify key mechanisms of pest regulation in polycultures.
They explain why there may be differences in mechanisms between cropping sys-
tems, and suggest plant assemblages which enhance regulatory effects and those
which do not, and under what management and agroecological circumstances.
According to these theories, a reduced insect pest incidence in polycultures may be
the result of increased predator and parasitoid abundance and efficiency, decreased
colonization and reproduction of pests, chemical repellency, masking and/or feeding
inhibition from nonhost plants, prevention of pest movement or immigration, and
optimum synchrony between pests and natural enemies (Andow, 1991).
A recently conducted, well-replicated experiment, where species diversity was
directly controlled in grassland systems, found that ecosystem productivity was
increased and that soil nutrients were utilized more completely when there was a
greater diversity of species, leading to lower leaching losses from the ecosystem
(Tilman et al., 1996). In agroecosystems, this same pattern applies to insects as
herbivore regulation increases with increasing plant species richness. Evidence sug-
gests that as plant diversity increases, pest damage tends to reach acceptable levels,
thus resulting in more stable crop yields (Figure 6). Apparently, the more diverse
the agroecosystem and the longer this diversity remains undisturbed, the more
internal links develop to promote greater insect stability. It is clear, however, that
the stability of the insect community depends not only on its trophic diversity but
on the actual density-dependence nature of the trophic levels (Southwood and Way,

Figure 6 Hypothetical trend of pest regulation or damage reduction as species richness
increases in agroecosystems. “X value” represents the level at which a functional
assemblage of species with natural control attributes is established.
1970). In other words, stability will depend on the precision of the response of any
particular trophic link to an increase in the population at a lower level. Thus, selective
diversity, rather than just a random collection of species, is crucial to achieve desired
pest regulation (Dempster and Coaker, 1974).
From a practical standpoint, it is easier to design insect manipulation strategies
in polycultures using the elements of the natural enemies hypothesis than those of
the resource concentration hypothesis. This is mainly because we cannot yet
identify the ecological situations or life history traits that make some pests sensitive
(i.e., their movement is affected by crop patterning) and others insensitive to
cropping patterns (Kareiva, 1986). Crop monocultures are difficult environments
in which to induce the efficient operation of beneficial insects because these systems
lack adequate resources for the effective performance of natural enemies, and
because of the disturbing cultural practices often utilized in such systems. Poly-
cultures already contain specific resources provided by plant diversity and are
usually not disturbed with pesticides (especially when managed by resource-poor
farmers who cannot afford high-input technology). They are also more amenable
to manipulation. In polycultures, the choice of a tall or short, early or late maturing,
flowering or nonflowering, legume or nonlegume companion crop can magnify or
decrease the effects of particular mixtures on specific pests (Vandermeer, 1989).
Thus, by replacing or adding the correct diversity to existing systems, it may be
possible to exert changes in habitat diversity that enhance natural enemy abundance
and effectiveness.

Figure 7 The effects of landscape fragmentation on the expected viability of natural enemy
populations in agroecosystems of varying scales and levels of artificialization.
PATTERNS OF LANDSCAPE STRUCTURE AND

INSECT BIODIVERSITY
An unfortunate trend accompanying the expansion of agricultural monocultures

is that it occurs at the expense of surrounding natural vegetation which serves to
add biodiversity to the landscape. One consequence of this trend is that the total
amount of habitat available for beneficial arthropods and natural enemies of pests
is decreasing at dramatic rates. The hypothetical impact of habitat fragmentation on
the survival of natural enemies in agroecosystems is depicted in Figure 7. The
implication of this habitat loss for the biological control of pests may be serious in
light of scientific data demonstrating increased abundance of insect pests in homo-
geneous agricultural landscapes (Altieri and Letourneau, 1982). Emerging data dem-
onstrate that there is enhancement of natural enemies and more effective biological
control where wild vegetation remains at field edges and in association with crops
(Altieri, 1994). These habitats may be more important as overwintering sites for
predators, or they may provide increased resources, such as pollen and nectar for
parasitoids and predators from flowering plants (Landis, 1994).
Hedgerows and other landscape features have received significant attention in
Europe regarding their effects on arthropod distribution and abundance in adjacent
crop fields (Fry, 1995). There is wide acceptance of the importance of wild vegetation
field margins as reservoirs of natural enemies of crop pests (van Emden, 1965).
Many studies have documented the movement of beneficial arthropods from margins
into crops, and higher biological control is usually observed in crop rows close to
wild vegetation edges than in rows in the center of the fields (Altieri, 1994).
In many cases, weeds and other natural vegetation around crop fields harbor
alternative hosts/prey for natural enemies, thus providing seasonal resources to

bridge gaps in the life cycles of entomophagous insects and crop pests (Altieri and
Whitcomb, 1979). A classic case is that of the egg parasitoid wasp, Anagrus epos,
whose effectiveness in regulating the grape leafhopper, Erythroneura elegantula,
was increased greatly in vineyards near areas invaded by wild blackberry (Rubus
sp.). This plant supports an alternative leafhopper (Dikrella cruentata) which breeds
in its leaves in winter (Doutt and Nakata, 1973). Recent studies show that French
prune orchards adjacent to vineyards provide overwintering refuges for Anagrus and
early benefits of parasitism are promoted in vineyards with prune trees planted
upwind from the vineyard.
Research in northern California showed a considerable amount of movement of
entomophagous insects from woodlands into adjacent apple orchards, with organi-
cally managed orchards exhibiting a higher rate of natural enemy colonization from
bordering woodlands than orchards sprayed with insecticides (Altieri and Schmidt,
1986). Several predators and parasites collected in the woodland edges were inter-
cepted at the orchard interfaces and later collected within the orchards, suggesting
that the development of apple orchard beneficial arthropod communities are influ-
enced by the type of surrounding natural vegetation.
Recent developments in temperate agriculture to encourage predators while
reducing pesticide applications have included use of beetle banks, flowering strips,
and conservation headlands. In Britain, several hundred potentially beneficial species
of predators and parasites may live in or by cereal crops. Most of these are killed
when the crops are sprayed to control pests. But if the field habitat is manipulated
to increase plant diversity, then the need for spraying pesticides can be greatly
reduced. When grass strips are constructed across large fields, then predatory beetles
proliferate and can get to the field centers, the regions where aphid populations are
greatest (Wratten, 1988). The cost of establishing a 400-m bank in a 20-ha field is
about $200, including cultivation, grass seed, and loss of crop. One aphid spray
costs $750 across the same field, plus the cost of yield reduction due to aphid
infestation.
Despite the above findings, no major efforts are under way in the world to
diversify agroecosystems at the landscape level with natural edges or windbreaks
composed of flowering species that act as insectary plants. Experiments of this sort
would fill an information gap on how changes in the physical and biodiversity layout
of agroecosystems would affect the distribution and abundance of the whole complex
community of pests and beneficial insects.
Determining the dispersal of insects in response to landscape vegetational diver-
sity and whether or not natural plant strips surrounding crop fields serve as a
movement corridor for beneficial arthropods in monocultures will have major impli-
cations for planning IPM at the landscape level. It is expected that corridors can
serve as a conduit for the dispersion of predators and parasites within agroecosys-
tems. Given the high edge-to-area ratio in the corridors, this feature is expected to
have a high degree of interaction with adjacent crops, thus providing protection
against insect pests within the area of influence of the corridor by allowing distri-
bution of natural enemies within a certain range of the field. By documenting the
effects of the corridor on arthropod distribution and abundance, then it may be

Figure 8 Effects of diversification schemes (field margines, windbreaks, etc.) at the landscape
level on agroecosystem functions, with special emphasis on the dynamics of arthro-
pods. (Modified after Fay, 1995.)
possible to determine the length, width, distance, and frequency of corridors needed
to maintain a level of functional biodiversity which will provide the necessary crop
protection without the need of pesticides. A system of corridors can also have positive
effects on the overall system by interrupting inoculum dispersion, serving as barriers
to insect pest movement, producing biomass for soil fertility, decreasing outflow of
silt and nutrients, and by modifying microclimate through interception of air currents
(Figure 8). The most important aspect is that corridor manipulation can be a crucial
first step in reintroducing biodiversity into large-scale monocultures, thus facilitating
the biological restructuring of agroecosystems for the conversion to agroecological
management.
CONCLUSION
Diversified cropping systems, such as those based on intercropping and agrofor-

estry or cover cropping of orchards, have been the target of much research recently.
This interest is largely based on the new emerging evidence that these systems are
more sustainable and more resource-conserving (Vandermeer, 1995). Much of these

attributes are connected to the higher levels of functional biodiversity associated
with complex farming systems. In fact, an increasing amount of data reported in the
literature documents the effects that plant diversity have on the regulation of insect
herbivore populations by favoring the abundance and efficacy of associated natural
enemies (Altieri, 1994). Several hypotheses are emerging postulating the mecha-
nisms explaining the relationships between plant species number and the stabilization
of agroecosystem processes including the buffering of populations (Tilman et al.,
1996). One aspect that is clear is that species composition is more important than
species numbers per se. The challenge is to identify the correct assemblages of
species that will provide through their biological synergisms key ecological services
such as nutrient cycling, biological pest control, and water and soil conservation.
The exploitation of these synergisms in real situations involves agroecosystem
design and management and requires an understanding of the numerous relationships
among plants, herbivores, and natural enemies. Clearly, the emphasis of this
approach is to help to restore natural control mechanisms through the addition of
selective biodiversity within and outside the crop field, through a whole array of
possible crop arrangements in time and space.
Data and practical experience indicate that it is possible to stabilize the insect
communities of agroecosystems by designing and constructing vegetational archi-
tectures which support populations of natural enemies or that have direct deterrent
effects on pest herbivores (Altieri, 1991). What is difficult is that each agricultural
situation must be assessed separately, since herbivore–enemy interactions will vary
significantly depending on insect species, location and size of the field, plant com-
position, the surrounding vegetation, and cultural management. One can only hope
to elucidate the ecological principles governing arthropod dynamics in complex
systems, but the biodiversity designs necessary to achieve herbivore regulation will
depend on the agroecological conditions and socioeconomic restrictions of each area.
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CHAPTER 6
Livestock and Biodiversity
Harvey W. Blackburn and Cornelis de Haan
CONTENTS
Introduction
Livestock Production Systems
Grazing Systems
Mixed Farming Systems
Industrial System
Livestock System Interaction with Biodiversity
The Plant Community
Wildlife Interactions
Promotion of Livestock and Biodiversity
Technologies
Policies
Conclusion
References
INTRODUCTION
The manner in which human populations utilize livestock influences biodiversity.

If livestock are concentrated too much, the competition between wildlife and live-
stock increases or livestock alter the physical environment making it unusable by
some animal or plant species. But evidence is growing that when livestock are used
in balance with environmental resources they can actually enhance habitat for wild-
life. Much of the driving force determining how people use livestock, and therefore
the impact of livestock on the environment and biodiversity, stems from issues of

Table 1 Regional Growth Estimates (%) for
Demand of Meat and Cereals from
1990 to 2020
Region Meat Cereal
World 60–93 49–65
Developed 17–18 19–33
Developing 123–206 68–91
Sub-Saharan 141–194 136–161
Latin America 76–105 53–77
West Asia and North Africa 104–157 74–100
Rest of Asia 148–255 64–85
human population growth and economic development. There are options that can
help mitigate the negative impacts of livestock and biodiversity, and those shall be
explored in this chapter. For discussion purposes in this chapter we consider bio-
diversity to mean not the total number of species present in a specific ecosystem,
but rather the presence of critical types of species which permit ecosystems to
appropriately function. Given this definition, our contention is that if markets and
policies are appropriate, then livestock can help preserve biodiversity.
Globally the demand for livestock products is increasing and will continue to
grow (IFPRI, 1995). IFPRI (1995) data in Table 1 present regional growth rates in
the demand for meat compared with cereals. This growth in consumption of livestock
products is being fueled by economic and population growth throughout the devel-
oping world. As livestock numbers grow, there are direct implications for the envi-
ronment and biodiversity as a subset of any specific environment. There has been
concern that livestock have had a detrimental impact on the environment. However,
we shall see that this image is often incorrect, as much is dependent upon the human
population pressure and how those pressures display themselves. In other words, as
human population pressures increase, people can use livestock in a manner which
is detrimental to biodiversity.
LIVESTOCK PRODUCTION SYSTEMS
There are three principal types of livestock production systems that interact with
biodiversity: grazing systems, mixed farming systems, and industrial systems. All
three are found globally. Because these systems are so diverse in structure and
environment, it is difficult to make generalized statements about their impact on
biodiversity.
Grazing Systems
Grazing systems are defined as animal agricultural systems which are exclusively
livestock and have little, if any, crop production grown in conjunction with the
grazing of livestock. In these systems, livestock obtain most of the feed from native
vegetation. These systems are the most variable and diverse of livestock production

systems because of their dependence upon natural vegetation which is controlled by
weather changes. Grazing systems tend to be closed systems, where animal waste
products (manure) are used by the system.
In arid rangelands, livestock mobility is key to successful maintenance of vibrant
livestock and wildlife populations. These regions have tended to be the most con-
troversial areas of livestock use. However, a series of studies does show that the
extent of environmental degradation has been exaggerated. Three different works
demonstrate this point. R. Mearns (unpublished data) concluded that abiotic factors
such as rainfall, rather than livestock density, determine long-term primary plant
production and vegetation cover. Tucker et al. (1991) demonstrated the resilience of
arid lands using satellite imagery. Their work showed movement of the southern
belt of the Sahara depends on rainfall and that the southern boundary of the Sahara
is moving north. This movement is occurring after the long droughts which occurred
in the 1970s and is contrary to how we normally view the resource base in this
region. de Haan et al. (1997) demonstrated that, in the Sahel, livestock productivity
in terms of meat production per hectare and per head has increased over the past 30
years. This type of long-term trend would be difficult to obtain if pastoralists
constantly degrade the environment they utilize.
Semiarid and subhumid rangelands are more static systems than arid zone range-
lands. Due to the higher rainfall levels, these areas contain larger amounts of plant
biomass and have considerable biodiversity. These areas also receive heavier live-
stock grazing and allow opportunities for mixed farming systems to expand. As a
result of growing human population pressures, these lands and the biodiversity they
contain may be of most concern. For example, data from Mali showed that land
degradation in the 600 to 800 mm rainfall area was significantly greater than in the
350 to 450 mm rainfall area. However, such a trend is by no means a global
phenomenon. As Milchunas and Lauenroth (1993) demonstrate, in the semiarid and
subhumid regions, moderate grazing had no impact on biomass production, species
composition, and root development.
Livestock grazing in tropical rain forests (the humid zone) has more than any-
thing else typified the negative effects of livestock development on the environment.
Table 2 provides estimates of the main causes of deforestation (Bruenig, 1991). In
the humid region, data on biodiversity losses in the rain forest are dramatic. Since
1950, the world has lost about 200 million ha of tropical forest, with the resulting
loss of unique plant and animal species. Forested areas of Central America have
declined from 29 to 19 million ha since 1950, although after 1990 the rate of
deforestation in Central America decreases. In the 1980s rain forest in Central
America disappeared at the rate of 30,000 ha/year; this had declined to 320,000 ha
Table 2 Estimated Causes of Deforestation (percent of

total deforestation)
Region Crops Livestock Forest Exploitation
South America 25 44 10
Asia 50–60 Negligible 20
Africa 70 Negligible 20

over the period 1990 through 1994. In South America, the deforestation rate in the
1980s was about 750,000 ha/year. It is not known whether or not this rate has
declined over the last years. In Brazil, about 70% of the deforested areas is converted
into ranch land.
Much of the deforested areas in Latin America went into ranching, after initially
being cropped. In Central America, pasture areas have increased from 3.5 million
to 9.5 million ha and cattle populations more than doubled. For example, in Central
America, livestock have increased from 4.2 million head in 1950 to 9.6 million in
1992 (Kaimonitz, 1995). In Asia and sub-Saharan Africa, the decline in the forest
area is mainly the result of crop expansion.
In the temperate zones, extensive grazing livestock are usually produced with
some harvested forage during the wintertime. Livestock produced in extensive
grazing systems interact most closely with biodiversity in the form of wildlife. In
both Australia and the U.S. these systems were heavily used in the late 19th century,
and as a result degradation of the resource base occurred. Heavy use during the last
century led to legislation controlling the use of the grazing resource and conse-
quently has promoted rangeland health and improved range condition. Key issues
driving how people utilize temperate ranges include fuel prices and privatization
(Central Asia), heavy levels of fertilization (Europe), and concern over riparian areas
(western U.S.).
Mixed Farming Systems
In mixed farming systems, crops and livestock are produced on the same
resource base. Globally this system produces the largest share of meat (54%) and
milk (90%). Throughout the developing world, mixed farming is the main agricul-
tural production system for smallholders. Developmentally, mixed farming systems
provide farmers with an opportunity to reduce financial risk and smooth out pro-
duction cycles. Farmers are able to take the highs and lows out of their production
cycles because they have the capacity to provide livestock with higher-quality
forages during winter or dry months of the year. In return, the sale of livestock
products helps finance inputs for the farming enterprise; in addition, livestock
provide traction for soil preparation. The mixed farming system is also a partially
closed system where the manure can be utilized on the farm to build soil fertility
while the milk and meat produced in the system flow out to urban markets. In many
respects mixed farming systems have the capacity to promote healthy ecosystems
and provide for economic development of the farmer, but, due to human population
pressure, poverty, and poor infrastructure, these systems can negatively impact
biodiversity and the environment at large.
Mixed farming systems tend to be transitional as livestock production shifts from
an extensive grazing system to an intensively managed industrial system. McIntire
et al. (1992) have documented the role population pressure has in integrating crop
and animal agriculture and in promoting the integration of crop–livestock systems.
They also discuss how further increases in population pressure drive farmers to
become more specialized, therefore, causing the decomposition of the mixed farming
system into more-intensive crop or livestock enterprises. Disaggregating livestock

and crop agriculture may result in lower levels of biodiversity, for by keeping the
two activities biodiversity can be promoted and may prevent the agricultural system
from becoming too brittle (Holling, 1995).
Globally mixed farm types and the way livestock are used are extremely diverse.
In Southeast Asia, for example, livestock and crop production is very intensive.
Cattle are used for draft purposes and in turn consume high proportions of crop
aftermath. In contrast, many mixed farming systems in temperate Organization for
Economic Cooperation and Development (OECD) countries had and have the poten-
tial for being balanced systems. These types of farms have the capacity to produce
various crops (e.g., maize) in a rotation with alfalfa, which in turn provides a forage
resource for ruminant livestock and helps to replenish soil nutrients extracted in the
production of cereal grains. Mixed farming and biodiversity interact at several crucial
levels. First is the interaction with wildlife which can be positive or negative. Second,
by replenishing soil fertility through manure application, the mixed farm can help
provide a viable environment for soil microflora and microfauna. In developed
countries there has been a tendency for farmers to focus on monoculture crop
production. From a plant and animal perspective this makes these systems more
brittle and exposed to major stresses. By maintaining livestock in these farming
systems there is an opportunity to keep these systems more robust and encourage
more fully the presence of various plants and animals.
Mixed farming systems can be broadly classified into those found in developed
and developing countries. Developing country mixed farming systems contain sev-
eral environmental issues which impact biodiversity. Soil erosion impacts both the
people and plant/animal biodiversity in the various mixed systems. Pimental et al.
(1995) estimated erosion rates of 30 to 40 ton/ha/year in some Asian, African, and
South American systems. Bojos and Casells (1995) determined that in Ethiopia soil
loses were 5 ton/ha/year in grazing lands used by crop–livestock farmers while
erosion rates on crop lands were 42 ton/ha/year. Here livestock may be critical in
maintaining soil fertility and soil organic matter levels. In Southeastern Asia adding
pig and ruminant manure together may contribute up to 35% of the soil organic
matter requirements, therefore providing an important source of organic matter. This
is a crucial contribution because it is the only avenue available for farmers to improve
soil organic matter (de Haan et al., 1997).
In developed countries, soil erosion and soil fertility are issues that impact biodi-
versity. In temperate zones soil losses of up to 15 ton/ha/year have been reported by
Pimental et al. (1995). Soil fertility is impacted more by overfertilization than a lack
of soil nutrients. Once soils become saturated with excess levels of nitrogen or phos-
phorus, these nutrients leach into above- and belowground water systems. Driving
much of this overfertilization is the ease of importing feed and inorganic fertilizer. By
importing these products into the mixed farming system, there becomes less of a need
to balance animal feed and cropping activities through rotation and fallow systems.
Industrial System
The industrial system can be the most capital intensive of the livestock production
systems. In general, it is a large concentration of livestock (particularly poultry and

swine), but can also include small-scale periurban production in developing coun-
tries. Industrial systems do not produce their own feed; rather it is imported into the
system from other locations within a country or, in some extreme cases, it is imported
from other regions of the world. Industrial systems can impact biodiversity locally
through the wastes they generate or off-site, where the feed is grown for use in the
industrial system.
The industrial system has a threefold effect on biodiversity through:
• Waste production and its effects on terrestrial and aquatic ecosystems. These effects
are often geographically confined to areas of high livestock densities. Eutrophica-
tion and destruction of habitats is a common phenomenon in parts of northeastern
Europe and the U.S. as well as in the densely populated areas of the developing
world, in particular Asia and, to a lesser extent, Latin America. Ammonia emission
leads to acidification of the environment and negatively affects ecosystem func-
tioning and biodiversity.
• Demand for concentrate feed and resulting changes in land use and cropping
intensity. The production of feed grains, in particular, adds additional stress on
biodiversity through habitat loss and damages in ecosystem functioning.
• The requirement for extremely uniform animals of similar genetic composition
contributes to within-breed erosion of domestic animal diversity.
But the industrial system has many advantages. First, the rapid development of
industrial pig and poultry systems helps reduce total feed requirements of the total
livestock sector to meet a given demand. Therefore, it may help to alleviate pressures
leading to deforestation and degradation of rangelands, such as is happening in parts
of Latin America and Asia, thus saving land and preserving biodiversity. Second,
the feed-saving technologies developed for this system do not have scale effects and
can be successfully transferred to mixed farming systems. The same holds true for
animal waste prevention and treatment technologies that have been developed fol-
lowing regulations applied mainly to the industrial system. Therefore, the demand-
driven industrial system generates a series of innovations that have spillover effects
on the sector as a whole.
LIVESTOCK SYSTEM INTERACTION WITH BIODIVERSITY
The Plant Community
Native plant communities naturally go through a series of successional changes,

from low to high to low plant diversities while in the process of obtaining a state
of climax (Clements, 1905). Grazing by livestock overlies this natural process.
That is, grazing intensity interacts with and can modify the rate at which plant
communities move toward climax. In addition, there is some evidence that indicates
that for grazing to effect plant communities significantly requires a combination
of grazing intensity and rainfall or fire events (Milchunas et al., 1988; Westoby et
al., 1989).

An important concept in determining the status of plant community health is
that of thresholds (Westoby et al., 1989). The threshold concept proposes that plant
communities under grazing pressure do not deteriorate in a linear fashion. Rather,
there is a series of levels which a plant community moves to when confronted with
a series of pressures. Thresholds separate these levels. Within a level, plant commu-
nities can fluctuate in terms of biomass production and species composition, and
recovery within a level is more easily accomplished (Archer et al., 1988). If grazing
pressure is relaxed prior to a critical level or threshold, plant community recovery
becomes less problematic.
Depending upon how livestock graze in a specific environment, biodiversity can
increase or decrease. Either heavy or light grazing can lead to a reduction in
biodiversity. Moderate grazing tends to promote patchiness of vegetation (CAST,
1996). Increased patchiness allows for diverse plant species to compete in a given
environment. Therefore, by moderately grazing native rangelands, plant communities
can be manipulated to maintain a desired level of plant diversity.
The semiarid and subhumid areas are some of the world’s most important
repositories of plant and animal biodiversity. For example, in Africa, Le Houerou
(1991) estimates that rangelands contain about 3500 plant species, having a signif-
icant role in ruminant nutrition.
For the subhumid savannas, weed invasion is a major problem threatening biodi-
versity, and the role of livestock is only secondary. For example, the grass Imperata
cylindrica in the Philippines and Indonesia now has infested more than 5 million
ha. Invasion with broad-leafed plants and shrubs is more common in the savannas
of Africa and the Americas.
There are a large number of cases that show that in well-balanced grazing
systems, especially those using multispecies, plant biodiversity increases. An exten-
sive review of grazing and production data of 236 sites worldwide, including many
sites in the semiarid zone, showed no difference in biomass production, species
composition, and root development in response to long-term grazing in the field
(Milchunas and Lauenroth, 1993).
Wildlife Interactions
There are a variety of ways in which livestock can interact with wildlife com-
munities. These include (Burkholder, 1952; Odum 1971; Mosley, 1994):
1. Neutralism, where neither species affects the other;

2. Direct interference or resource use competition, where both species inhibit each
other;
3. Amensalism, where one species is inhibited and the other not affected;
4. Predation, where one species inhibits another by direct attack;
5. Commensalism, where one species is benefited by the presence of another but there
is no impact on the second species;
6. Protocooperation, in which the interaction between species is favorable to both
species but the association is not obligatory; and

7. Mutualism, the interaction is favorable to both species and the association is
obligatory.
The extent to which the interaction between livestock and wildlife is neutral, nega-
tive, or positive is dependent upon how domestic livestock are managed in specific
situations. Severson and Urness (1994) identify four ways livestock can be used to
modify species that can, in turn, develop habitats that are favored by certain wildlife
species. Such a modification is achieved by altering the composition of vegetation,
increasing the productivity of selected species, increasing nutritive quality of forage,
and increasing diversity of habitats by altering plant structure.
Riparian health is an important issue driving the monitoring and use of public
grazing lands. However, it is often overlooked that any species of wildlife or
livestock can overgraze these critical areas. A key example of such a situation exists
today in Yellowstone National Park (YNP), the crown jewel of the U.S. national
park system. It has recently been demonstrated that elk are severely overgrazing
riparian areas in YNP. In a study comparing riparian areas in YNP and on the summer
range of the U.S. Sheep Experiment Station (approximately 30 miles from YNP),
it was shown that grazing of sheep had a more beneficial impact on riparian health,
as measured by willow populations, a key indicator species (Figure 1). Furthermore,
as a result of healthier willow communities on the Sheep Station, beaver populations
are also in better condition. This work demonstrates that any grazing animal can
cause environmental instability and/or degradation and that by using an appropriate
livestock species environmental health can be maintained or increased (Kay and
Walker, 1997).
Another key aspect which determines the type of wildlife–livestock interaction
is diet preferences. Different types of wildlife and livestock prefer different plant
types. For example, cattle select more grass in their diet than sheep which choose
a combination of grass, forbs, and browse. The same type of diet selection patterns
are evident in wildlife. Murray and Illius (1996) cite examples in the Serengeti of
how small-bodied species, such as the Thompson’s gazelle, are more selective
Figure 1 Impact of elk and sheep grazing on willow communities: a measure of ecosystem
health and herbivore grazing.

grazers than larger animals, such as topi and buffalo. By having diets which do not
overlap helps maintain a broad diversity of plants. They further discuss the fact that
grazing pressure in the Serengeti increases the overall spectrum of resource avail-
ability to animal communities. By cropping and trampling the tall grasses, larger
ungulates increase the range and sward structures providing room for a greater
variety of ungulate species.
The interaction between wildlife and livestock in ecosystems can be complex.
First, there is increasing evidence of a grazing complementarity between wildlife
and livestock. As shown by Schwartz and Ellis (1981), the dietary “overlap” between
most wildlife species and livestock is rather limited. Mwangi and Zulberti (1985)
and Western and Pearl (1989) showed that the combination of livestock raising and
wildlife management resulted in an equal or better species wealth than any of these
activities done individually. Furthermore, in national parks in Kenya such as
Amboseli, where livestock is not permitted, biodiversity is decreasing, with an
increase in unpalatable plant species and bush encroachment (W. K. Ottichilo,
unpublilshed data). On the other hand, the same author points out that there are
many degraded areas in Kenya due to combined wildlife–livestock pressure.
The driving forces leading to losses in animal biodiversity are habitat destruc-
tion, species introduction, and hunting (World Resources Institute, 1994). Habitat
destruction is playing an important role in the developing world, especially in the
subhumid savannas. In Africa, road construction and human immigration from the
drier areas leads to habitat destruction of the vectors of African sleeping sickness.
In turn, this lifts the protection of wildlife, which is tolerant to the disease. Inter-
national agencies, including the World Bank, have also financed extensive vector
clearance campaigns in West Africa. Traditionally, these campaigns used a combi-
nation of hand and aerial insecticide spraying, initially with organochlorines, to
eradicate the tsetse fly. Nagel (1993) argues that pesticides from this period are still
notable in some African birds.
Since the mid 1980s, compounds with shorter residual effect, such as synthetic
pyrethroids, have been used. These second generation compounds caused substantial
initial damage to the flora and fauna, but permanent effects were not observed with
single spraying, for example, in the World Bank–funded tsetse clearance project on
the Adamaoua Plateau of Cameroon (P. Muller, unpublished data). The permanent
damage and high residue levels reported from this project came from repeated
spraying in the border areas. Bush encroachment resulting from inappropriate graz-
ing management was the most serious environmental damage. Land-use plans have
often been advocated as the essential elements of tsetse clearance, and international
financiers have made the preparation of such plans conditional to the financing of
the eradication campaigns. However, the experience with the enforcement of such
land-use plans has been dismal, as local authorities lacked the authority and means
for their enforcement. A critical issue concerning these zones is that traditional land
tenure practices have been even less robust in these more humid areas than in the
drier areas. This lack of strong traditional tenure practices has been felt in sub-
Saharan Africa after tsetse clearance operations, and as a result a rather anarchic
settlement pattern developed.

In addition, hunting and culling of wildlife was encouraged in the past, because
wildlife was thought to be a reservoir of diseases, such as rinderpest and malignant
catarrhal fever, and carriers of disease, such as East Coast fever and trypanosomiasis
(Grootenhuis et al., 1991), and competition for scarce grazing resources. However,
the control of the above-mentioned diseases has improved considerably and there is
a much better understanding of which particular species harbor specific diseases.
The costs and returns from wildlife, compared with livestock and agricultural
production, are highly variable. At the level of the national economy, the opportunity
cost of wildlife biodiversity conservation in protected areas, in terms of forgone
livestock and agricultural production, seems to outweigh the income from tourism
and forestry generated by these protected areas. For example, in Kenya, Norton-
Griffiths and Southey (1995) estimated the forgone livestock and agriculture from
the parks at U.S.$ 203 million, while the revenue from these parks amounts to only
U.S.$ 42 million. On the other hand, Engelbrecht and van der Walt (1993) estimated
that the Kruger Park in South Africa contributed more than U.S.$ 110 million/year
in tourism vs. a forgone production of only U.S.$ 6 million. At household levels,
the comparative profitability of wildlife and livestock raising varies greatly, accord-
ing to the ecological conditions and wildlife use (meat, trophy hunting, tourism).
Overall, under present conditions of niche markets for game meat or tourism,
wildlife ranching seems financially more attractive, though. For the communal
areas, wildlife cannot provide the multiple functions of producing milk for subsis-
tence and providing traction, fertilizer, and investment that livestock can. Without
any doubt, the combination of wildlife and livestock is the most appropriate under
those conditions.
A recent World Bank study gives comparative income levels from wildlife and
livestock production on ranches for four African countries. In Ghana, investments
in cattle had an economic rate of return (ERR) of close to zero, whereas private
wildlife ranching had an ERR of about 8. In Kenya, financial return on investment
(FRR) from game ranching was estimated at 7 to 12% vs. about 6 to 8% from
livestock ranching. In Namibia, the study gave a wide range of net returns varying
from 0 to 0.28 rands/ha under livestock, to 0.28 to 1.50 rands/ha under wildlife. In
Zimbabwe, the FRR for livestock was about 2%, compared with 10% for wildlife.
Economic returns per hectare were higher for ranching than for wildlife. All wildlife
enterprises benefited from the special niche market, either through higher meat
prices, or through revenue from tourism or trophy hunting (Bojos, 1996).
In fostering sustainable wildlife–livestock integration on communal areas, insti-
tutional constraints play a big role. Traditionally, there has been a rigid centralized
and regulatory attitude of public institutions in the protection of wild animals. This
was especially the case in East Africa, where wildlife management was typically
organized by central administrations in a rather military fashion. There was no benefit
sharing with the local population, whereas wildlife causes high financial cost to the
local population because of crop damage and livestock loss due to predators and
diseases. This has led to antagonistic reactions from many herding and farming
communities. In addition, the interdiction, still in effect in many countries, of sport
hunting and consumption precludes benefits from wildlife to be realized as an
important part of the potential benefits (W. K. Ottichilo, unpublished data).

PROMOTION OF LIVESTOCK AND BIODIVERSITY
Literature cited in this chapter demonstrates that livestock can have either a
positive or negative impact on biodiversity. The driving forces are human population
growth, economic development, and how societies within a country value their
natural resource base and, in particular, biodiversity. Given that livestock can have
either a positive or negative impact on biodiversity and the use of livestock is based
upon larger policy and economic issues, what are the avenues that promote a positive
symbiotic relationship between livestock and plant and other animal species? Is there
a group of technologies and policies that can be implemented or practiced which
can promote an agenda involving economic growth and maintenance of biodiversity?
The use and development of such technologies and/or policies can be critical to
sustaining a rich array of biodiversity.
Technologies
Increasingly, the scientific community is determining that there are a variety of

technologies that can promote economic growth of the livestock sector and encourage
or enhance biodiversity. For example, in Southeast Asia as well as in the North
American Pacific northwest, there is a growing appreciation that sheep can be used
in forestry systems (rubber or timber, respectively) to control undesirable plant
species. The significance of this approach is that it reduces the amount of herbicide
that has to be applied. The development of the above practices demonstrates that
there is capacity to develop technologies and practices that successfully blend live-
stock production with biodiversity issues.
The following list presents ways which could facilitate the development of
technologies that both increase livestock productivity and promote biodiversity.
Generalized technologies needed for grazing systems include
• Better quantification of the global economic costs and benefits of livestock use and
development with conserving biodiversity;
• Methodologies for identification of appropriate indicators that provide reliable
information on plant and animal trends;
• Utilization of mixed species grazing (cattle, sheep, goats with wildlife) at appro-
priate levels — mixed species grazing has the advantage of increasing plant com-
munity diversity while concurrently improving the habitat of wild ungulates, but
there is a need to develop methodologies which can more accurately determine
optimal combinations of domestic and wild ungulates;
• Design of sustainable drought preparedness plans in arid and semiarid areas which
not only take into account the needs of livestock and their owners but the impact
their activities will have on plant and wildlife communities;
• Development and use of breeds of livestock which are well suited to the environ-
ment in which they are expected to perform, such as indigenous breeds.
Generalized technologies needed for mixed farming and industrial systems

include

• Improved soil cover through use of alternative crops for mulching;
• Improved feed production and quality to reduce the pressure on grazing areas and
improved internal nutrient transfers;
• Reduced nutrient losses from manure and improved efficiency of application (both
actions promote biodiversity either through the contribution of manure to soil
organic matter or by preventing the over application);
• Improved feed formulation which better balances animal dietary needs and reduces
nitrogen excretion;
• Improved animal management through better matching the nutrient needs and use
of genotypes which best match the environment;
• Development and utilization of precision agriculture to determine the amount, type,
and benefit of inputs needed.
Policies
Both livestock sector and biodiversity issues are affected by broader policy issues
that a country may deem necessary to implement. For example, a country may decide
to import cereal grains for livestock during times of drought rather than support a
rational destocking program which would lead to more productive livestock sectors
and support a quicker range recovery after the drought has ended. Biodiversity is
compromised by the value the society of a country places upon its natural resource
base. In other words, demand for food and economic growth can overwhelm any
concern for the environment. Cheap food policies and a desire to achieve self-
sufficiency, particularly with cereals, have been important factors in determining
livestock sector growth and biodiversity conservation. Furthermore, overvalued
exchange rates in sub-Saharan Africa and Latin America have favored importation
of cheap food from the industrialized world, thus competing against local production
and providing few incentives for local producers to intensify into mixed crop–live-
stock systems and to practice soil conservation.
The following list of policy options could contribute to a better blending of
livestock development and conservation of biodiversity:
• Eliminate overvalued exchange rates which favor importation of cheap food from
the industrialized world, thus competing against local production and providing
no incentives for local producers to intensify into mixed crop–livestock systems
and practice soil conservation.
• Strengthen land tenure security, especially in the rain-fed mixed farming systems
of the developing world that will provide an incentive for investment in long-term
soil fertility improvements, such as the use of inorganic fertilizers and the use of
green manure and leguminous fodder crops in the crop rotation.
• In dry areas, improve infrastructure, roads, and markets to facilitate movement of
goods and services especially in drought. Carefully introduce water development
so that vegetation is not negatively impacted and aquifers are not depleted.
• Develop institutional capacity that blends biodiversity and livestock needs and
provides a basis for analyzing and evaluating economic and environmental needs.
• Develop more effective benefit-sharing mechanisms for communities practicing
livestock production in a manner which helps promote biodiversity.

• Phase out subsidies on feed, fertilizers, and mechanization to promote a tighter
integration of crop–livestock systems and remove opportunities to overfertilize.
• Tax inorganic fertilizer, set maximum application limits, and regulate time of
application to reduce leaching and volatilization.
CONCLUSION
As research continues on the relationship between livestock and the environment,

and specifically on biodiversity, there is a growing awareness that livestock can have
positive impacts on biodiversity as well as a negative relationship. Driving the
livestock–biodiversity relationship are the pressures human populations place on the
natural resource base. Negative impacts on plant and wildlife communities occur
when livestock either outcompete wildlife (usually through high stock rates) or
overgraze an area for long periods of time, thus causing permanent damage to a
plant community. However, appropriate stocking rates have been shown to improve
biodiversity in both the plant and animal communities involved. CAST (1996) states
that wildlife habitat has generally improved in conjunction with improving range
conditions in the western U.S.
Due to the increased needs of human populations on a global scale, agricultural
pressures, including livestock, will continue to mount and potentially deplete some
portion of biodiversity. But as de Haan et al. (1997) discuss, it is imperative to have
appropriately formulated policies which lessen the negative relationships between
livestock and biodiversity and strengthen the positive aspects of the relationship.
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CHAPTER 7
Managing for Biodiversity of Rangelands
Neil E. West
CONTENTS
Introduction
Definitions
A Case Study in Biodiversity
Sagebrush Steppe
Location, Ownership, and Land Uses
Climate
Primary Producers
Consumers
Decomposers and Nutrient Cycling
Interactions among Plants, Animals, and Humans
Preservation of Relatively Unaltered Ecosystems
Alteration of Existing Heavily Grazed Stands
Rehabilitation of Burned Sagebrush Steppe
Regional Considerations
Guidelines for a New Style of Rangeland Management Sensitive to
Biodiversity
Conclusions
References
INTRODUCTION
Under accelerating extinctions within world biota and increasing invasion of

exotics, consequent to the expansion of human populations and their increased

demands for space devoted to producing their immediate needs and aspirations
(Vitousek et al., 1997), numerous environmental interest groups have clamored for
more consideration of natural biotic wealth of all kinds. In the past these efforts
focused on creation of reserves. Activists, however, now realize that increasing the
size of existing reserves and demarcation of new ones will not conserve all the
biodiversity many would like. Furthermore, changing climates mean that fixed
boundary reserves will not guarantee that suitable habitat will be available for
organisms to migrate to (Harte et al., 1992). Conservation biologists (e.g., Noss and
Cooperrider, 1994) are thus shifting some of their attention to nonreserve lands of
all kinds and attempting to alter land-use policies such that biodiversity is provided
for over a larger fraction of the Earth.
Rangelands, where native biota intermingle with humans and their domestic
livestock, involve a huge fraction of the Earth’s surface (about 70% by the estimate
of Holechek et al., 1989). Increasing conflict between graziers and conservation
biologists seems inevitable, especially in the developed world where people have at
least the short-term luxury of considering wildlife and other amenities over produc-
tion of food and fiber. The fact that the wildlife are owned by most states, whereas
most habitats are owned by individuals or local communities (Cumming, 1993), is
the major reason for biodiversity issues providing clashes between private rights and
public values, particularly on publicly owned lands.
DEFINITIONS
Before we go further, we need to define some critical terms. First, one needs to
realize that biodiversity entails many different things to different interest groups
(West, 1995). To some, it is mainly genetic material. To others, it is taxonomic
richness, usually species, of biota within plots or more abstract communities and
landscapes. To still others, it is properly functioning ecosystems, including indige-
nous human cultures living in sustainable ways. All these views are legitimate and
have to be respected in democratic societies.
Even though scientists of various kinds are pushing broadened views of biodi-
versity, the public activists are, as reflected in legislation, budgets, and activity,
favoring the charismatic megafauna, the warm, fuzzy, and appealing organisms,
particularly the vertebrates, not the little things that run the world (Wilson, 1987).
Administration of the Endangered Species Act (ESA), the strongest environmental
law in the U.S., currently only impacts what can be done to listed species and their
habitats, including activities on privately held lands and waters.
It is becoming obvious that far more than scientific information is involved in
what is being done about biodiversity. Stances about biodiversity inevitably involve
one’s personal and professional ethics (Coufal, 1997). Thus, this is a topic that will
inevitably cause philosophical reflection, as well as scientific and managerial action.
The second term deserving further definition is rangelands. Some prefer a strictly
use-oriented definition. In that sense, rangelands are agroecosystems since they are
all lands with self-sown vegetation used for livestock grazing. That is the oldest

definition that still prevails in developing countries. This traditional definition also
applies to a wide array of ecosystem types where livestock grazing has and could
occur, including recently cut forests, tundras, and marshes. The majority of range-
lands, however, occur where grasslands, shrub steppes, deserts, woodlands, or savan-
nas prevail, in other words, most of the untilled or undeveloped western U.S. (about
70% of the area). Rangeland managers and scientists are thus more familiar with
drier and less fertile systems than most foresters, wildlife biologists, and agrono-
mists. Whereas most of such lands were recently seen primarily as sources of food
and fiber, in developed countries many of them are being increasingly dedicated to
sustaining other values that are now prized more highly in industrialized societies.
We thus have to contrast how rangeland biodiversity is being considered in the
developed compared with the developing world.
My focus here will be on the drier parts of the world where self-sown vegetation
is managed extensively based on ecological principles. Agronomic principles rarely
apply to these lands: the costs of attempting to till, seed, fertilize, treat with pesti-
cides, and use other means of strong manipulative control to enhance production of
food and fiber rarely justify their expenditure because plant responses are funda-
mentally low due to meager precipitation, salty, steep, and rocky soils, etc. The
previous lack of such treatments is the major reason that rangelands are now seen
as valuable repositories of biodiversity. That is, most of these rangelands have not
yet been simplified and homogenized by intensive agricultural activities (Matson et
al., 1997). There are some important exceptions, however, such as the Conservation
Reserve Lands (Allen, 1995), which are former croplands that could become range-
lands and/or wildlife reserves, depending on Congress’ budget setting.
A CASE STUDY IN BIODIVERSITY
Sagebrush Steppe
A thorough review of all aspects of biodiversity in all kinds of rangelands around

the world would be impossible for several reasons. First of all, not all aspects of
biodiversity have been thoroughly studied in all kinds of rangelands. The genetics
of even dominant plants and vertebrates, and anything about invertebrates, microbes,
ecosystem functions, and feedbacks, have rarely been studied. Second, even the
information that does exist cannot all be summarized in the space available here.
Therefore, what I have chosen to do is exemplify how biodiversity issues interact
with science and policy in one ecosystem type (sagebrush steppe) well known to
the author. I will bring in ideas and experimental results from other contexts as well
and discuss how they might apply to sagebrush steppe. In that way I can give a
more-focused introduction to the topic at hand.
Shrub steppes are ecosystems with organisms and life-forms of both deserts and
grasslands. Although, on average, they are drier than most grasslands and wetter
than deserts, the variation in climate is high (coefficients of variation in total annual
precipitation usually exceed 30%). Thus, some years have grasslandlike climate

whereas other years are desertlike. This climatic variation is probably the main
reason for the mix of grassland and desert life-forms in making up shrub steppes.
Another result of the high climatic variation is the inherently low stability of these
systems under disturbance (Archer and Smeins, 1991).
Because the environmental conditions of the sagebrush steppe are harsh and
highly variable over time and space, the dominant organisms are few and widely
distributed. This belies the probable high degree of intraspecific ecotypic and genetic
variation, which has barely been studied. Once these patterns are understood, vari-
ations in autecological and ecophysical responses and synecological interactions will
be more comprehendible.
Location, Ownership, and Land Uses
Sagebrush steppe occurs wherever there is or once was vegetation with shared
dominance by sagebrushes (woody Artemisia spp.) and bunchgrasses (West and
Young, 1998). This system occurs mostly in the lowlands of the northern part of the
Intermountain West. Sagebrush steppe once occupied about 45 × 106 ha there (West
and Young, 1998). About 20% of this ecosystem type passed into private ownership
with the Euroamerican settling of the West (Yorks and McMullen, 1980). The remain-
ing 80% is managed by various agencies of the U.S. and state governments. This
circumstance makes the management of these lands much more difficult than those
under private ownership. Many interest groups, including those championing bio-
diversity, can and do politically influence management policies on these public lands.
About half of the original sagebrush steppe area now in private ownership has
been converted to either dryland or irrigated agriculture over the past 150 years. The
approximately 90% remaining untilled lacks irrigation water or is too steep, rocky,
or shallow soiled for annual cultivation. The dominant historical uses of these
wildlands by human societies have been first hunting and gathering and then live-
stock grazing.
Climate
The prevailing climate in sagebrush steppe is temperate, semiarid (mean annual

precipitation of 20 to 40 cm) and continental (cool, wet winters and springs and
warm, drier summers and autumns). Mean annual temperatures range from 4 to
10°C. Winters are cold enough so that snow packs of 50 to 100 cm are common.
Snowmelt is usually gradual and thus most of the moisture therein becomes stored
at depth in the soil. Native plant growth occurs largely from April to July, the only
part of the year when both temperatures and soil moisture are favorable. Summer
precipitation is rarely enough to carry herbaceous plant growth throughout the
summer. Early fall precipitation is not dependable and by October temperatures are
usually too cool to allow much regreening of grasses (West and Young, 1998).

Primary Producers
The major woody dominants here are woody Artemisia, collectively known as
the sagebrushes. These are shrubs derived from progenitors which came from Eurasia
over the Bering Land Bridge and have subsequently radiated into about 13 species
(McArthur, 1983). Furthermore, the major species, Artemisia tridentata (big sage-
brush), has at least five relatively easily recognizable subspecies that should be used
in separating out different ecological sites (McArthur, 1983).
The sizes and degrees of dominance of the sagebrush species vary greatly with
both site and disturbance history. Sagebrush density is generally greater, but height
lower, on more xeric sites. Sagebrush also increases in abundance following exces-
sive livestock grazing in the spring (West and Young, 1998). Livestock grazing also
reduces the chance of fires by removal of fine fuels in the interspaces connecting
the clumps of shrubs. Fire formerly kept the sagebrush steppe more frequently burned
(60 to 110 year return interval) (Whisenant, 1990) and less dominated by sagebrush
because most species of sagebrush do not resprout after fire, but have to regenerate
from seed (Blaisdell et al., 1982).
Even when sagebrush is dominant, a moderate number of other plant species are
found associated with it. On relict (naturally ungrazed by livestock) sites in central
Washington, Daubenmire (1970) found an average of 20 vascular plant species in
1000-m plots. Tisdale et al. (1965) found a range of 13 to 24 vascular plant species
on three relict stands in southern Idaho. Zamora and Tueller (1974) found a total of
54 vascular plant species in a set of 39 late seral stands in the mountains of northern
Nevada. Mueggler (1982) found between 24 and 41 vascular plant species in a set
of 68 0.05-ha lightly grazed macroplots in sagebrush steppe of western Montana.
The vertical and horizontal plant community structures are remarkably similar in
all relatively undisturbed examples of this ecosystem type. The shrub layer reaches
approximately 0.5 to 1.0 m in height. The shrubs have a cover of about 10 to 80%,
depending on site and successional status. The grass and forb stratum reaches to about
30 to 40 cm during the growing season. Herbaceous cover also varies widely depend-
ing on site and successional status. On relict sites, the sum of cover values usually
exceeds 80%, and can approach 200% on the most mesic sites (Daubenmire, 1970).
The herbaceous life-forms most prevalent on relict sites are hemicryptophytes
(Daubenmire, 1975). The proportion of therophytes increases markedly with distur-
bance. The proportion of geophytes is around 20%. A microphytic crust dominated
by mosses, lichens, and algae is commonly found where litter from perennials is
not excessive (West, 1990). Sagebrushes have both fibrous roots that can draw water
and nutrients near the surface and a taproot that can function from deep in the soil
profile. Near the end of the growing season for grasses, sagebrushes nocturnally
water from more than 90 cm and excrete it in the upper part of the soil profile at
night (Caldwell and Richards, 1990). This hydraulic can help the grasses stay active
longer than possible on their own.
Perennial grasses associated with Artemisia vary greatly throughout the region.
The C bunchgrasses (Agropyron spicatum, Festuca idahoensis, Stipa spp., Sitanion
hystrix, Poa spp.) dominate the herbaceous layer in the north and western parts of
the type. C sod grasses (e.g., Agropyron smithii, Hilaria jamesii) become more

common in the south and east where more growing season precipitation occurs
(West, 1979).
Total aboveground standing crop phytomass within the sagebrush steppe type
varies between about 2000 to 12,000 kg/ha, depending on site differences, successional
status, and age of the brush (West, 1983). Litter standing crops are about one half
the live nonwoody material (West, 1985). Belowground phytomass is similar in
magnitude to that aboveground. Annual net aboveground primary production varies
between about 100 and 1500 kg/ha, depending on site, successional status, stand
age, and preceding climatic conditions (Passey et al., 1982).
Plant ecologists have long assumed that communities that are floristically richer
stabilize primary production in the face of variable climate (Chapin et al., 1997).
Indeed, Passey et al. (1982) in their discussion of long-term data gathered from
ungrazed sagebrush steppe relicts conclude that each year brings both unique dom-
inance–diversity and production relationships. They attribute this to differing phe-
nologies, rooting patterns, and green leaf persistence. Harper and Climer (1985)
reanalyzed the Passey et al. (1982) data set and concluded that variation in plant
community production was more positively related with floristic richness than either
average precipitation or precipitation of a given year. Tilman et al. (1996) have
shown that greater species richness in tall grass prairie leads to greater production
during drought than in more depauperate stands created by adding nutrients.
Any landscape within which sagebrush steppe is the matrix is a patchwork of
stands of differing species composition and shrub or other growth form dominance.
The mix of plant species and growth forms is dependent on ecological site potential
and time since particular disturbances. Fires, grazing by both native and introduced
vertebrates and invertebrates, as well as unusual climatic events such as deep soil
freezing before snowpack accumulation and unusually heavy precipitation and con-
sequent soil anoxia, all contribute to resetting the successional clock (West and
Young, 1998). Livestock grazing on these rangelands usually takes place in large
paddocks with only one or a few watering points. The parts most distant from water
thus are less grazed and of higher seral status (Hosten and West, 1996). This creates
a patchwork of differing seral statuses across the landscape (Laycock et al., 1996).
Consumers
The native vertebrates using this ecosystem type are a mixture of grassland and
desert species. Maser et al. (1984) grouped the vertebrates of sagebrush steppe in
southeastern Oregon into 16 life-forms and related them to vegetation structure and
other features of habitat. The vertebrate community is more diverse when the
vegetation has the greatest structural diversity (Parmenter and MacMahon, 1983).
Neither shrub-dominated nor grass-dominated situations favor as many different
kinds of vertebrates as do the mixtures. A few such as voles (Microtus montanus)
can influence the structure by girdling the shrubs (Mueggler, 1967; Parmenter et
al., 1987).
Over 1000 species of insects have been observed on a sagebrush–grass site in
southern Idaho (Bohart and Knowlton, 1976). Wiens et al. (1991) recently identified
76 taxa of invertebrates on sagebrush alone in central Oregon. Relatively little is

known about the habitat preferences, trophic relationships, and other aspects of the
roles of invertebrates in this ecosystem type. Only a few — thrips, webworms,
grasshoppers, cicadas, aphids, and coccids (Kamm et al., 1978; West, 1983) — are
known to be irruptive and visibly alter vegetation structure.
Decomposers and Nutrient Cycling
Very little is known about microbes and the decomposition process in this
ecosystem type. Initial studies of the nitrogen (West and Skujins, 1978) and phos-
phorus (West et al., 1984a) cycles showed that available forms of these elements
may limit plant production in wetter than average years. Allelochemics from sage-
brush and the high C:N ratios of its litter may inhibit some decomposition and
nitrogen-cycling processes, perhaps indirectly strengthening sagebrush dominance
in this ecosystem type (West and Young, 1998). Changes in litter quality can lead
to degradation of soil organic matter in such systems (Lesica and DeLuca, 1996).
Global environmental changes may produce some unexpected interactions among
plants, soil microbes, and soil degradation (West et al., 1994).
Interactions among Plants, Animals, and Humans
The pristine sagebrush steppe evolved with large browsers (megafauna), most
of which had disappeared by about 12,000 years ago (Mehringer and Wigand, 1990;
Burkhardt, 1996). The loss of the megafauna is inextricably linked to simultaneous
increases in human hunting and climatic warming (Grayson, 1991). Remaining
graminivores were few in the pre-European system (Mack and Thompson, 1982;
Harper, 1986). The small populations of aboriginal hunters and gatherers of the mid-
Holocene probably influenced the vegetation largely by burning. It took European
colonization to change drastically the native vegetation and the wildlife habitat it
provides (Young, 1989).
The pre-European era livestock grazing capacity, when shrubs were fewer and
grasses more prevalent, was estimated to be 0.83 animal unit months (AUM)/ha
(McArdle and Costello, 1936). Because sagebrushes are usually unpalatable to
livestock, whereas herbs are palatable, uncontrolled livestock use led to a decline
of herbs and increase in brush. Carrying capacities declined to an average of 0.27
AUM/ha in the 1930s (McArdle and Costello, 1936), but had improved slightly to
0.31 AUM/ha by 1970 (Forest-Range Task Force, 1972).
Livestock populations built up rapidly near the end of the 19th century. Griffiths
(1902) judged that the grazing capacity of these rangelands had been exceeded by
1900. Hull (1976) examined historical documents and concluded that major losses
of native perennial grasses and expansion of shrubs took only 10 to 15 years after
a site was first grazed by livestock.
The native grasses are extremely palatable, especially when green. They die
easily when grazed heavily in the spring (Miller et al., 1994). In addition, they rarely
produce good seed crops (Young, 1989).
The only time the grasses and forbs have an advantage over brush is when sites
are burned. However, on the sites with heavy historical livestock use, both remaining

native herbaceous perennials and their seed reserves have been greatly diminished
(Hassan and West, 1986). In addition to tall, thicker sagebrush, grazing-induced
freeing of space and resources gave opportunities for the invasion of aggressive
Eurasian plants. The advent of introduced winter annual grasses, notably Bromus
tectorum in the 1890s (Mack, 1981), and the continuous, fine, and early-drying fuels
they provide has led to seasonally earlier, more frequent (less than 5 years), and
larger fires (Whisenant, 1990). After repeated fires, combined with unrestricted
grazing, any remaining native vegetation becomes easily replaced by other, even
more noxious introduced annuals, such as medusahead (Taeniatherum caput-medu-
sae), knapweeds (Centaurea spp., Acroptilion spp.), and yellow star thistle (Centau-
rea solstitialis). The result has been a considerable decrease in plant species structural
and floristic diversity, average forage production, and nutritional value to vertebrates
(Billings, 1990; Whisenant, 1990). This simplification of self-sown vegetation results
in much more frequent bare ground and accelerated wind and water erosion (Hinds
and Sauer, 1974). Variability in plant production goes up several orders of magnitude
after replacement with annuals (Rickard and Vaughn, 1988).
Wildlife responds dramatically to these changes in vegetation structure (Maser
et al., 1984). For instance, the pigmy rabbit (Brachylagus idahoensis) is a threatened
species that prefers the tallest, densest stands of Basin big sagebrush. Sites occupied
by this plant have been widely converted to intensive agriculture. Thus, the range
of this sensitive animal has been reduced and its abundance greatly diminished.
Another native herbivore of special interest in the sagebrush steppe is the sage
grouse (Centrocercus urophosianus). This is a large galliform with a unique digestive
system that has coevolved with Artemisia. The mature birds survive the less hospi-
table times of the year by eating the twigs of sagebrushes, especially the low
sagebrushes found on windswept ridges. There are, however, other requirements
during other parts of their life cycle. During March and April, the males gather on
open areas without brush (called leks) and display themselves to the females. Only
about half of the males survive raptor predation and intraspecific fighting during this
about 2-week mating period. The females fly to the most productive interfluvial
areas to nest and raise the chicks. For the first 6 weeks of life, the young birds
require a high protein diet made up of insects and forb buds. These are most abundant
in fresh burns and in riparian corridors.
Sage grouse were very abundant in the region when Europeans first arrived and
have remained abundant enough to be an important game bird until recent decades.
Unfortunately, they are now being considered for placement on the endangered lists
in several Intermountain states. Wildlife and conservation biologists find it tempting
to single out the range livestock industry for causing this problem. However, sheep,
which prefer forbs over other types of forage, were much more abundant on these
rangelands up to about 1960, but have since declined to a tiny fraction of their former
abundance. Sheep do, however, eat some sagebrush, particularly in the fall and
winter. The amount of time cattle are permitted on public lands of the sagebrush
steppe has also been declining since about 1964, well before sage grouse populations
crashed. The amount of perennial cover on much remaining sagebrush steppe has
been increasing of late because of reduced livestock grazing and more effective fire
control. There is now probably more sagebrush than necessary for optimum sage

grouse use in most portions of the sagebrush steppe. Several other possible influences
have also been increasing of late, such as vehicular access and nonhuman predators.
Coyotes, foxes, skunk, racoons, corvids (jays, magpies, crows, and ravens), and
raptors (eagles, hawks, and owls) have all been increasing because of less shooting
and pesticide use and could be taking more eggs and chicks, as well as adults. The
thickened brush could be making predator stalking and capture easier.
Because of passage of laws such as the ESA and National Forest Management
Act, the interests of wildlife, particularly the rare, endangered, and threatened ver-
tebrates, can take precedence over optimal livestock grazing on publicly owned
rangelands in the U.S. This is the reason that the U.S. Forest Service and Bureau
of Land Management currently strives to leave about 15 to 20% of the mature
sagebrush cover intact across the landscape rather than burning or using herbicides
to reach the 100% kill they once strived for in the 1940s and 1950s when the nation
demanded more red meat.
There has already been a vast replacement of native plant species by Eurasian
plant invaders in sagebrush steppes. More is expected, especially if global warming
materializes. Controlling fires entirely is an impossibility. Reductions or even com-
plete removal of livestock will not result in a rapid return to the vegetation that
occurred before European colonization (Miller et al., 1994). Sheep, grazed during
the fall, because they utilize some sagebrushes and can do little damage to the
herbaceous understory during that time of year, can actually enhance floristic rich-
ness (Bork et al., 1998).
Our major means of obtaining greater dependability of forage production and
soil protection on severely degraded sagebrush steppe sites, while at the same time
reducing the chance of fire, has been to plant Eurasian wheatgrasses and ryegrasses
(Asay, 1987). However, this can only be done easily on relatively level sites with
deep, largely rock-free soils. Environmental and archaeological interest groups have
recently stopped these procedures, however. Environmentalists object to using any
introduced species, regardless of their ability to grow rapidly and protect the soil.
Archaeologists object to the physical disturbances to archaeological objects and
strata. Native species have been repeatedly tried in plantings, but rarely grow early
and rapidly enough to outcompete the introduced annuals. Because environmentalists
have prevailed, public land managers are no longer daily involved in proactive
management or ecosystem repair here.
Let us now turn to other possible ways to conserve remaining community
diversity, alter existing stands, or rehabilitate degraded sagebrush steppe stands.
Figure 1 will be used to guide the following discussion. This figure is a state-and-
transition model (Laycock, 1995) thought to accommodate better our current under-
standing of degradation and successional processes in sagebrush steppe than the
simpler, linear models of the past with one end point (the climax).
Preservation of Relatively Unaltered Ecosystems
Pristine, relictual areas (State I in Figure 1) no longer exist nor are probably
recoverable. The reasons for this view are

Figure 1 State-and-transition model of successional change in sagebrush steppe.
1. Humans (indigenous peoples) are no longer hunting, gathering, and burning these
areas. The previous fire regimes are no longer in place and as the vegetation changes
in response to less frequent fires, the hydrologic and nutrient cycles are being
altered, as is the habitat for numerous animals and microbes.
2. The present climate is warmer and drier than the cooler, wetter Little Ice Age
climate which prevailed up to about 1890. Thus, only heat- and drought-tolerant
species may thrive now under global warming.

3. Atmospheric CO has increased about 20% during the past century, altering the
competitive balances in this vegetation as well as changing the nutritional qualities
of the phytomass and litter (Polley, 1997).
4. About 15% of the flora is now new to the region.
Since we can reverse none of these influences, at least in the short term, we should
learn to live with what remains and manage it toward the desired plant communities
we choose for each circumstance.
There are, however, some remnants of these landscapes that have escaped direct
human influences. These relics exist because they have no surface water, are sur-
rounded by difficult topography, or protected in special-use areas (e.g., military
reservations). I place these in State II of Figure 1. Tisdale et al. (1965) describe an
example. I estimate that less than 1% of the sagebrush steppe that remains has
avoided the direct impact of any livestock. Even these relicts are, however, incom-
plete because of lack of indigenous humans and lengthened fire frequencies. Relicts
are influenced by air pollutants, climatic change, and invasion by exotics (Passey et
al., 1982). Most of the existing late seral sagebrush steppe (State II in Figure 1) has
had light livestock use. Even light livestock use puts inordinate pressure on a few
highly palatable species (ice cream plants), partially explaining the lack of a return
arrow from State I to State II.
In some places, feral horses and burros now put considerable pressure on such
rangelands, but are protected by federal law on most public lands. I estimate that
about 20% of the remaining sagebrush steppe is in State II.
The perceived will of a majority of Americans now is to identify these remaining
State II areas, especially those on public lands, and protect them from being devel-
oped. Some advocate all such areas be reserved (Kerr, 1994), whereas others (Bock
et al., 1993) propose that 25% have livestock excluded. Rose et al. (personal com-
munication) have, however, recently demonstrated that lightly grazed sagebrush
steppe has higher species richness than adjacent exclosures dating to 1937. Others
propose restoration efforts to bring further-degraded systems back to States I or II
(Dobson et al., 1997). State II areas serve as the parts catalogue for restoration
efforts. The Gap Analysis Program (GAP) of the U.S. Fish and Wildlife Service
(Scott et al., 1993) and the various natural heritage programs initiated by the Nature
Conservancy are well under way to put these views in action. These efforts are,
however, not without attack from both political and scientific groups (Machlis et al.,
1994; Short and Hestbeck, 1995).
I expect to see physical modifications to enhance production of food and fiber
(formerly called range improvements) to be more spatially limited than in the past
because such actions on public lands or with public monies require environmental
assessments or impact statements and thus public scrutiny and debate. The remaining
relatively unaltered areas on public lands will probably be consciously protected to
provide the later seral condition patches necessary to hold a broader spectrum of all
species, and meet the special requirements for some featured species such as sage
grouse and pigmy rabbit (Call and Maser, 1985). Of special concern are other
sagebrush bird obligates that are also apparently declining: sagebrush sparrow

(Amphispiza belli), sage thrasher (Oreoscoptes montanus), and Brewer’s sparrow
(Spizella breveri).
Rangeland managers in the past strove to reduce the limitations of the land for
producing livestock. These limitations were mainly topography, forage availability,
and water. For example, trails were constructed into areas where topographic breaks
limited livestock access. Natural water was supplemented by development of springs,
building stock tanks and small dams, drilling wells, piping and hauling water. Fences
were constructed and salt distributed to control livestock movement and institute
grazing management systems (e.g., rest–rotation grazing). All these improvements
were designed to distribute livestock utilization more uniformly across the land, gain
greater efficiency of food and fiber production, and divert livestock from the espe-
cially sensitive riparian areas (Elmore and Kauffman, 1994; Laycock et al., 1996).
The net result has been progressively more widespread intensive use of a landscape
that has become partially tamed from the wild. These assumptions need to be
reexamined in the light of biodiversity concerns. Let us continue our consideration
of these relationships in the sagebrush steppe.
Alteration of Existing Heavily Grazed Stands
Because livestock grazing of native sagebrush steppe usually avoids the unpal-
atable forages, particularly woody species, they are freed from competition and
dominance becomes concentrated in the few woody plants on areas with a history
of heavy livestock grazing (T2), but not recent fire (State III, Figure 1). About 30%
of this ecosystem type is estimated to exist currently in this state. Most of these
stands can stay stagnated for decades (Rice and Westoby, 1978; West et al., 1984b;
Sneva et al., 1984; Winward, 1991). The dense, competitive stands of excess sage-
brush prevent the herbaceous species from recovering. Such brush-choked stands
are usually chosen by both livestock and wildlife managers for manipulation to
diversify vegetation structure. This enhances it for livestock or native animals in
spots, concentrating livestock use, reducing their pressure elsewhere, while simul-
taneously advantaging some wildlife species through vegetation modifications via
grazing systems, prescribed burning, brush beating, or chaining (T3). For example,
sheep grazing in the fall, because they consume more sagebrush then (Bork et al.,
1998), can be used to obtain a reversal from State III to State II. Prescribed burning
(Harniss and Murray, 1973) can also be applied to stands with sufficient remnant
populations of native herbs to quickly recover following brush kill. Rest from
livestock use, such as with a rest–rotation grazing system or winter only use (Mosely,
1996), will often allow a slower return to State II from State III. Reduction of brush
also enhances water yields (Sturges, 1977), and some seeps, springs, and streams
reappear. When phenoxy herbicides are used alone (Evans et al., 1979) (T4) or in
conjunction with fire, the community becomes dominated with native grass (State
IV, Figure 1) because the chemicals impact all broad-leafed species. The conversion
only slowly returns (T6) to State II with judicious grazing and a secondary treatment
with prescribed burning. About 5% of the remaining sagebrush steppe is now esti-
mated to be in State IV. This is a short-lived state, especially under heavy grazing
(T5). Mueggler (1982) found enhanced alpha diversity in moderately grazed sage-

brush steppe communities in western Montana following prescribed fire, 2,4-D, and
brush-beating treatments. Summer fires can damage some of the grasses (Young,
1983), but encourage the resprouting rabbitbrushes (Chrysothamnus spp.) and horse-
brushes (Tetradymia spp.) (Anderson et al., 1996).
If accelerated soil erosion does not ensue and the fundamental potential of the
site does not change, then State III can be maintained or managed toward States II
or IV. However, as herbaceous plants and litter in the interspaces between perennials
are reduced, soil aggregate stability declines, infiltration of precipitation diminishes,
overland flow increases, and soil erosion frequently increases (Blackburn et al.,
1992). When a probable threshold of use is exceeded, the site can irreversibly change
to one of lesser potential. This explains the dashed line and downward arrows below
States III and V as the only believable transitions. This is where the syndrome of
desertification is most evident. All the former states can be dealt with via soft energy
management approaches. Once this threshold is exceeded, however, subsequent
management requires expensive, risky, hard energy solutions. Unfortunately, it is
often easier to get political attention after major damage has been done rather than
getting budgets and personnel to plan, monitor, and tweak the higher-condition,
more-natural systems at opportune times.
The desertified sites with thickened brush have largely introduced annuals in
their understory. I estimate that State V comprises about 30% of the current sagebrush
steppe. Reduction or removal of livestock only hastens further degradation from
State V because livestock remove part of the fuel load and thus reduce the chance
of fire destroying the sagebrush and the spots of soil it protects.
If insufficient amounts of native herbs remain on sagebrush steppe, the usual
land management agency response has previously been to replace them mechanically
(T7) with introduced wheatgrasses and ryegrasses, especially crested wheatgrass
(Asay, 1987). This has been done because the introduced perennial grasses are much
more easily established than the native grasses and they grow quickly to provide
more forage with a higher nutritional plane. The introduced perennial grass stands
are also much more tolerant of subsequent heavy livestock use and have lasted for
many decades (Johnson, 1986). There are some long-range concerns, however (Les-
ica and DeLuca, 1996), because the introduced perennial grasses suppress the return
of natives and richer plant species assemblages. Some large treatment areas have
monocultures of Eurasian perennial grasses prevailing (State VI, Figure 1). I estimate
about 5% of the original sagebrush steppe has already been transformed to State VI.
Wildlife biologists have noted declines in the numbers of birds (Olson, 1974;
Reynolds and Trost, 1979; 1981), small mammals (Reynolds and Trost, 1980), and
large reptiles (Reynolds, 1979) on such seedings of introduced grasses. It should be
noted, however, that such studies present a worst-case scenario because samples
came from the center of large treatments. Provision for increased diversity near
edges (Thomas et al., 1979) is not usually mentioned in such studies. Present-day
more-sensitized planners would provide for optimum edge effect and patchiness
(McEwen and DeWeese, 1987). When society makes the investment in repairing
severely damaged sagebrush steppe, creating perennial grass–dominated pastures of
much greater productivity of species palatable to livestock (T7), this should com-
pensate for livestock reductions and other management restrictions on lands where

States II, III, and IV (Figure 1) predominate. Because introduced grass pastures take
heavy degrees of utilization in the spring, the native shrub steppe can support fall
and winter grazing with less impact, especially on the native herbaceous perennials.
Introduced perennial grass plantings in the sagebrush steppe region, especially
if grazed by livestock, will eventually experience shrub reinvasion (T8 to State VII)
largely in response to intensity of livestock grazing. I estimate that about 5% of the
remaining sagebrush steppe region is currently represented by shrub-reinvaded intro-
duced wheatgrass/ryegrass pastures (State VII). Not all brush is now eliminated by
re-treatment (T9). Herbicide use on public lands in the Pacific Northwest has been
suspended by judicial decree. Prescribed burning of the coarser, introduced grasses
is difficult and leaves patches where the shrubs prevail. There are, therefore, chances
to enhance edge effects in large areas that were formerly homogenized. As in the
untilled native areas, we could enhance wildlife habitat by providing a mix of
successional stages or stand conditions, providing both cover and forage for either
featured species or total species richness (Maser et al., 1984). For example, some
success has been attained in creating alternate leks for sage grouse breeding follow-
ing disturbances of development (Eng et al., 1979).
Rehabilitation of Burned Sagebrush Steppe
Despite greatly increased attention to fire prevention and control, much of the
most-depauperized sagebrush steppe (State V) has been burned (T10) at least once
during the past three decades and is now dominated by introduced annuals, mainly
grasses such as cheatgrass and medusahead (State VIII, Figure 1). The Bureau of
Land Management (M. Pellant, personal communication) estimates that about 3
million acres of public lands in Idaho, Utah, Oregon, and Nevada are now dom-
inated by cheatgrass and medusahead. I estimate that about 25% of the total
original sagebrush steppe has made this transition (T10, T11). Because of their short
stature, restricted nutritional characteristics (short period of aboveground gree-
ness), and greater susceptibility to recurring fires than sagebrush steppe, such areas
are undesirable from all viewpoints. Without nutritional supplementation, livestock
can graze State VIII only during the short, early spring plant-growing season
(winter use is possible in the lower elevation areas near the Columbia River;
Mosely, 1996). Only the most generalist animals, such as the introduced chukars
(Alectoris chukar), horned larks (Eremophila alpestris), grasshoppers, and deer
mice (Peromyscus maniculatus) seem to thrive on the annual grasslands (Maser
et al., 1984). When such areas burn in early summer, soils are bared to wind and
water erosion during the convectional storms of summer. The consequent needs
for revegetation after fire are increasing while the budgets of federal land man-
agement agencies decline and pressure from environmentalists against active man-
agement increases.
Land dominated by annuals may provide fair watershed protection during years
without fire and actually appears to be more productive of total plant tissues than
the original sagebrush–native perennial grass and forb combination (Rickard and
Vaughan, 1988). This is likely, however, to be only a temporary situation based on
the priming effect of decomposing litter (Lesica and DeLuca, 1996) and the miner-

alization of nutrients from the enormous belowground necromass of the original
system. When these reserves of nutrients and soil organic matter are finally respired
away, the annual grasslands are likely to become much less productive. Similar
transitions happened in the Middle East several millennia ago (Zohary, 1973). Many
other, more noxious weeds from that region could find their way here, and we could
witness a downward spiral of further degradation (T12).
Rather than allowing the annual grasslands derived from former sagebrush steppe
(State VIII, Figure 1) to remain and the land to degrade further, some land managers
are attempting to intervene. A joint U.S. Forest Service, Bureau of Land Manage-
ment, Agricultural Research Service, and University of Idaho program is under way
to reduce these threats (Pellant, 1990). The most notable component of this effort
is the green-striping program most evident in southern Idaho. The basic approach
is to begin breaking up the now vast stretches of cheatgrass and other annual
dominance that have developed as fires have become earlier, larger, and more
frequent (Whisenant, 1990). Land managers are attempting to break the area into
smaller, burnable units, especially nearer to cities and towns. The approaches used
thus far include planting strips of vegetation that stay green (and thus wetter and
less burnable) longer than cheatgrass.
Although the introduced wheatgrasses and ryegrasses do stay green longer and
burn less readily, because of coarser aboveground structure, they are not native and
thus are rejected by some interest groups. Because the genetic biodiversity of the
native plants is so primitively understood, the best that can be done is to gather such
seed locally and plant it on comparable sites. Such seed sources are undependable,
however; thus a root-sprouting big sagebrush is seen as a potentially better keystone
species to put back in this area. A few sagebrushes may actually help sustain
perennial grasses by harboring the predators on black grass bugs (Labops spp.)
(Haws, 1987). Furthermore, total plant community production can be enhanced
(Harniss and Murray, 1973) because sagebrushes help trap blowing snow (Sturges,
1977) and scattered sagebrushes moderate temperatures (Pierson and Wight, 1991),
benefit the reestablishment of native herbs, and protect them from excessive utili-
zation (Winward, 1991). Sagebrushes also harbor mycorrhizal fungi (Wicklow-
Howard, 1989), which helps them extract nutrients from deep in the soil and recycle
them to the surface through litter production (Mack, 1977; West, 1991).
Whether or not we can accomplish restoration of sagebrush steppe (T13, between
States V and III in Figure 1) is highly questionable. Even where money is less
limiting and topsoil is stockpiled on coal strip mines, early results are only partially
encouraging (Hatton and West, 1987). We will have to know much more about how
sagebrush steppe ecosystems are structured and function and obtain vast budgets
and more trained personnel before such efforts are routinely successful. It is cheaper
and more feasible to foster good stewardship of land having late seral vegetation
(manage while in States I, II, III, or IV of Figure 1) rather than rely on restoration
efforts after degradation has taken place (States V, VI, VII, and VIII of Figure 1).

Regional Considerations
Because biodiversity issues in sagebrush steppe are interconnected to multiple

impacts and other ecosystem types over the entire region, the federal management
agencies are attempting to address them in a holistic fashion. An important example
of this is the proposal for ecosystem management in the interior Columbia Basin
(Haynes et al., 1996). The documents generated (Quigley et al., 1997) appear to
favor restoration practices. Environmentalists (e.g., Belsky, 1997), however, perceive
little change in livestock grazing practices and intend to test the process judicially.
GUIDELINES FOR A NEW STYLE OF RANGELAND MANAGEMENT

SENSITIVE TO BIODIVERSITY
Recent happenings in the Interior Columbia Basin are symptomatic of the start
of a new era in land management. When human populations were lower and demands
on resources were less, we could encourage development without much concern for
other species or equity to the future. It is becoming obvious now that more consid-
eration for present neighbors and future generations must be consciously given.
Environmental impacts no longer have only local consequences. Biodiversity can
be viewed as a natural treasure and as having a role in the maintenance, cleansing,
and repair of ecosystems at local to global scales (Chapin et al., 1997). Development
plans of the U.S. Agency for International Development and the World Bank now
require consideration of biodiversity within their environmental impact sections.
We are seeing enhanced efforts to inventory, monitor, and zone with biodiversity
in mind. National, regional, and local rankings of organisms and system rarity and
endangerment (e.g., the GAP analysis, Scott et al., 1993) are leading to plans to
create core reserves, buffer, corridor, multiple-use, and intensive-use zones. Some
graziers and other rangeland consumptive users are bound to be either displaced or
have their activities altered by these designations. The consequences could be com-
plete removal of livestock in some places of special sensitivity. In most other
situations, more thoughtful and careful pastoralism can complement conservation
(Friedel, 1994). Areas too small and dispersed to be managed efficiently within a
reserve could instead be managed by the permittee on public rangelands. Such areas
are called Excised Management Units in Australia (Morton et al., 1995). Where
livestock use is critical only at certain times, Restricted Use Units may be designated.
Where such designations cause economic hardship, land trades or subsidization may
help ease the transition. It is becoming clear that no modern government or non-
governmental entity can afford the expense of buyouts of increasingly greater blocks
of reserves. Furthermore, unless reserves are well managed, they can be just as
deleterious to the conservation of biodiversity as have been exploitative pastoral
systems. Such unmanaged areas can quickly become havens for predators, feral
animals, and noxious weeds (Friedel and James, 1995).
In cases where restoration is being attempted, it seems only right that displaced
graziers be employed to stay on the land and actively work to heal it. After all, these

are people who best understand the local environment. Their children should be
assisted in training for other jobs and professions.
Not all healing of degraded rangeland ecosystems requires complete displace-
ment of livestock grazing. Fleischner (1994) and Noss and Cooperrider (1994, Table
7.1) provide a comprehensive discussion of the negative ways management of
rangelands and attendant activities (e.g., irrigating winter fodder, predator control,
etc.) influence biodiversity. Brussard et al. (1994; 1995) and Brown and McDonald
(1995) point out the imbalances of Fleischer’s and Noss and Cooperrider’s presen-
tations. To help further balance those discussions (see Laycock et al., 1996), I wish
to add some positive aspects of the interactions of rangeland livestock husbandry
and biodiversity.
First, full-time ranching provides daily contact with the land at all seasons and
thus provides experience and a degree of attention to the land that occasional field
visits by agency personnel and intermittently interested environmentalists can never
replace. Indeed, the Nature Conservancy is calling on such full-time ranchers to
manage some of their properties actively with continued livestock grazing, yet with
enhanced sensitivity to biodiversity. The Nature Conservancy realizes that simply
buying up key properties and eliminating direct human influence is not a viable
way to conserve biodiversity on a grander scale. It realizes that humans are part
of the ecosystem and that it could not purchase and preserve all the desirable
properties anyway. Instead, it understands that encouraging management in eco-
nomically, as well as ecologically, sustainable ways is the long-term answer to
holding on to the maximum biodiversity across the rangelands of the western U.S.
The conservancy intends to lead by developing examples that neighboring ranchers
will emulate. It has already established worthy examples in places such as Red
Canyon Ranch, Wyoming.
Conservation biologists often mistakenly assume that, because mismanaged live-
stock have done much damage to rangelands in the past (a fact that even livestock-
oriented scientists don’t deny, e.g., Pieper, 1994), simply removing them perma-
nently will automatically result in the return to similitudes of a romanticized pre-
Columbian Eden. In many ways, Euroamericans have enhanced biodiversity by their
activities on rangelands (Johnson and Mayeaux, 1992). So many aspects of envi-
ronment and biota have and are currently undergoing change that it is fruitless for
us ever to expect equilibrial scenarios henceforth (Botkin, 1990; Pimm, 1991; Allen
and Hoekstra, 1992; Vitousek et al., 1997). The only rational choice is to monitor
and adjust through adaptive resource management (Kessler et al., 1992).
First, trade-offs must be made between maximum production of livestock and
the best possible wildlife habitat, watershed, and soil protection under the ecosystem
management philosophy (Kessler et al., 1992). No matter what the manager does
or does not do, habitat of some species will be enhanced and that of others simul-
taneous diminished (West, 1993). For instance, in a recent rangeland study in Aus-
tralia (James et al., 1997), about one quarter of the biota was disadvantaged by
livestock grazing, about half was neutral to grazing, and about one quarter was
increased. Humans have to make the choices of what is favored. Grazing by livestock
can be advantageous or disadvantageous to wildlife and other land uses depending

on species, uses and values, and their ecosystem context (West and Whitford, 1995).
Retention of total species richness, featured species, and stenotypic species cannot
usually be simultaneously maximized on the same small piece of land. Such objec-
tives have to be managed for on landscape and regional bases (Friedel and James,
1995), thus inevitably involving many landowners and institutions.
Midseral and even early-seral conditions are not detrimental to all wildlife
species. Some species require these conditions to complete their life cycle (West,
1993). Maser et al. (1984) found that the greatest species numbers and highest
population levels of most featured vertebrate species of sagebrush steppe are in
midseral condition. Thus, maintenance of disturbances to create early to midseral
conditions is desirable on at least some parts of a landscape. This is one reason why
total removal of livestock from public rangelands would not necessarily lead to
optimal habitat for either featured species or total species richness. Furthermore,
prohibiting livestock grazing would lead to more fine herbaceous fuels and thus
hotter, more frequent, and larger fires and eventual takeover by introduced annual
plants in sagebrush steppe (Figure 1) as well as many other kinds of adjacent
rangelands.
Domestic livestock management directly affects wildlife in two major ways: (1)
consumption of forage that could be used by wildlife and (2) alteration of vegetation
as it influences escape and thermal and protective cover (Noss and Cooperrider,
1994). Most animal species are more adapted to gross vegetation structure for
thermal and hiding cover than they are to particular plant species for food (Dealy
et al., 1981). Structural diversity of rangeland vegetation also relates positively to
wildlife species richness, except if the mosaic is on a scale too small to meet the
home range needs of species that require large blocks of uniform vegetation. For
example, optimum spacing between stands of big sagebrush and crested wheatgrass
for black-tailed jackrabbits (Lepus californica) requires that the wheatgrass openings
be no more than 600 m across because rabbit use of wheatgrass occurs mostly within
300 m of the type edge (Westoby and Wagner, 1973).
Domestic livestock management indirectly affects wildlife by (1) human and
livestock presence, (2) fencing, (3) salting, (4) water developments, (5) roads, (6)
trails, (7) predator control and other physical and chemical manipulations, such as
prescribed burning, chaining, cabling, root plowing, brush beating, reseeding, and
herbicidal application. The latter treatments usually simplify and homogenize habitat
structure, but mosaics and edge can be increased with planning and plant species
richness enhanced by interplanting in areas with large expanses of currently homo-
geneous vegetation.
Better planning and management could result in retention of livestock and their
use as tools for constructive improvement of wildlife habitat and watersheds. Deseret
Ranch in northern Utah is an example of a commercially viable operation that has
derived income from both consumable and nonconsumable wildlife while increasing
both livestock use and stabilizing range condition (Wolfe et al., 1997). Roads can
be closed and off-road vehicles prohibited, especially at critical times. Scattered
trees could either be retained or planted to provide shade and storm cover for both
livestock and wildlife, simultaneously enhancing their overall distribution.

Use of electronic sensors on livestock and thoughtful placement of invisible
electronic boundaries offer promise to foregoing building more fences and possibly
even to removing the existing ones eventually (Fay et al., 1989). We should recog-
nize, however, that fences serve as perch posts for some birds (Graul, 1980), and,
thus, unintended impacts of fence removal and pole line installation could ensue.
Some feel that there is promise in either selecting domestic animals that naturally
spend less time in sensitive areas (e.g., riparian zones) or training them to avoid
such areas through adversive conditioning. Indeed, some progress has been recently
made in doing just that (Howery et al., 1998).
Some water sources can be completely or selectively closed off to favor certain
species. Development of naturally occurring springs and seeps through installation
of perforated pipe and water troughs at a distance from the water source has made
more water available for drinking by both ungulates and birds. Unfortunately, the
natural wetlands around the original springs have often been highly altered. We can,
however, pipe out the overflow and create new fenced-out wetlands to replace those
altered (Kindschy, 1978).
We could even fertilize certain portions of some plant communities to increase
and freshen (make more palatable) some areas to draw animals to them to enhance
fire control, strutting, feeding, and nesting grounds. Guzzlers (artificial water catch-
ments) can be built in areas with limited free water for drinking by both wildlife
and livestock.
CONCLUSIONS
I have demonstrated that application of some knowledge, logic, planning, sen-

sitivity, and compromise could allow us to continue using most rangelands for
traditional values as well as provide for preservation and even enhancement of
biodiversity. Because of their rarity, desirability to research, and in guiding manage-
ment, most areas that have escaped livestock use thus far should probably be pro-
tected. This will provide maximum landscape diversity, reference areas for moni-
toring and basic research, and materials for restoration efforts. For the much larger
fraction of the rangelands that have had, and continue to have, extensive use, but
are still dominated by native plant species, we should continue to apply our increas-
ing knowledge of individual species responses, community dynamics, and ecosystem
feedbacks in devising low-input ways to direct succession toward desired sustainable
outcomes. Prescribed burning, behavioral modification of animals, and improved
ways of distributing animals without fences should be further developed. If some
compromise is deemed possible, livestock grazing systems compatible with wildlife,
recreational use, watershed and soil protection, as well as biodiversity, can be
devised. It is much cheaper and satisfying to prevent such seminatural areas from
slipping over the brink of irreversible trends toward desertification than trying to
rehabilitate or restore areas that have already been seriously degraded. We should
not let the possibility of artificial recovery prevent us from concentrating on trying
to develop sustainable use strategies for the rangelands still relatively intact.

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CHAPTER 8
Agroforestry for Biodiversity in

Farming Systems
Roger R. B. Leakey
CONTENTS
Introduction
Sustainable Production
Agroforestry
Agroforestry and the Diversification of Agroecosystems
Development of Multistrata Agroforests
Biodiverse Agroecosystems
Forest Patches, Biogeographical Islands, and Agroforestry
Domestication of Trees for Timber and Nontimber Forest Products
Commercialization
Tree Domestication in Progress
Conclusions
References
Abstract — Agroforestry can be used to diversify and intensify farming systems

through the integration of indigenous trees producing marketable timber and nontim-
ber forest products and is described in terms of an agroecological succession, in
which climax agroforests are biodiverse, highly productive, and profitable. The role
of biodiversity in agroecosystem function is one of the keystones of sustainability.
Complex agroforests that combine profitability with biodiversity are presented as a
model worthy of expansion. However, little is known ecologically about how best to
integrate agroforestry into the landscape, or to what extent agroforestry can be used
to link forest patches and expand biogeographical islands.

Tree domestication is one way to diversify and intensify agroforestry systems and to
make them profitable. A wise domestication strategy for indigenous trees will involve
the capture and maximization of intraspecific genetic diversity and so benefit both
production and the environment.
INTRODUCTION
The numbers of plant and animal species on Earth represents only 0.1% of those
that have existed since life appeared on this planet, the other 99.9% being already
extinct as a result of five episodes of mass extinction over geological time (Leakey
and Lewin, 1996) and the current period of extinctions (30,000/year against a
background of 0.25/year) arising from our population growth and lifestyle. Land-
use changes associated with colonization have been the major cause of these species
losses over the last few centuries. Agriculture, which has been described as the
“engine of economic growth” because of its powerful role in facilitating and stim-
ulating growth of other sectors of the economy (Mellor et al., 1987), started as a
subsistence activity 8000 years ago. The early subsistence systems are generally
considered to have been sustainable, while large-scale, capital-intensive, modern
agriculture has traded innate sustainability for chemical and other inputs and is
characterized by deforestation and a decrease in the overall numbers of associated
wild plants and animals to favor the growth of the planted crop. Typically, agriculture
systems and forestry plantations are monocultures of staple food or tree crops with
the almost total disappearance of the biodiversity and spatial complexity of natural
ecosystems. Characteristically, these monocultures are also based on the few plant
species that have been domesticated (Leakey and Tomich, 1998) and which also
have a narrow genetic base. In recent years, these developments have given rise to
concerns about deforestation, the loss of biodiversity, and the sustainability of our
lifestyle and, particularly, the crucial food production systems that are essential to
prevent famine and malnutrition in the tropics.
SUSTAINABLE PRODUCTION
Many of the means to increased productivity and profitability are now perceived
by society as carrying too high a cost in social disruption, human inequity, and
environmental degradation. The problem in trying to address this is how to define
and quantify sustainability. Izac and Swift (1994) developed an operational frame-
work to assess sustainability based on the premise that “a cropping system is
sustainable if it has an acceptable level of production of harvestable yield which
shows a non-declining trend from cropping cycle to cropping cycle over the long
term.” Their framework is based on the assessment of key ecological and economic
parameters at the field, farming system, and village/catchment scales and the concept
that a sustainable system never reaches threshold levels of irreversibility and that it
achieves a sufficient level of economic efficiency and social welfare. One of the

requirements identified by Izac and Swift (1994) is that the by-products (soil and
water quality, biological diversity, etc.) of agricultural activities must not disrupt the
biological functions of the system to the extent that the capacity of the system to
absorb these disruptions is surpassed.
Sustainability thus involves a symbiosis between the properties of the ecosystem
and the management activities that results in nondeclining and relatively stable
outcomes. Izac and Swift (1994) consider that the key to this symbiosis lies in the
assumed positive relationship of agroecosystem function to biodiversity and com-
plexity. Biodiversity therefore is a keystone in sustainability, and its loss has been
one of the common outcomes of agricultural intensification (Figure 1).
Agroforestry, through the replenishment of soil fertility and the domestication
of indigenous trees producing marketable forest products, has been proposed as one
way of diversifying and intensifying agroecosystems in a way that is beneficial to
the environment and can maintain and perhaps enhance biodiversity (Sanchez and
Leakey, 1998; Sanchez et al., 1998). In its Medium Term Plan 1998–2000, the
International Centre for Research in Agroforestry (ICRAF, 1997) foresees that agro-
forestry can contribute to human welfare and environmental resilience, with
improved systems providing:
1. Tree products that both increase food and nutritional security and generate cash
income for poverty alleviation and
2. Services that support and enhance ecosystem function (Figure 2).
The relevant services of trees are those that increase the crop yields (nitrogen
fixation, increased soil organic matter, nutrient cycling, soil conservation, etc.), create
environmental resilience (niche diversification, food web complexity, carbon seques-
tration, reduced greenhouse gas emissions, etc.), and provide social benefits (bound-
ary delineation, shade, etc.). Of these, the least is known about the ways in which
trees enhance the environment, although the body of information is increasing (see
Ingram, 1990; Swift et al., 1996).
AGROFORESTRY
Agroforestry, where it has been practiced traditionally, such as in the damar

agroforests of Sumatra and Jungle Rubber on Kalimantan (Michon and de Foresta,
1996) and in the home gardens of Sri Lanka (Jacob and Alles, 1987), Nigeria (Okafor
and Fernandes, 1987), and Tanzania (Fernandes et al. 1984), is a mixed and often
apparently haphazard polyculture of indigenous trees and crops that form a complex,
multistrata system somewhat like a natural forest. Interestingly, recent findings in
the damar agroforests of Sumatra show that these complex multistrata agroforests
contain over 50% of all the regional pool of resident tropical forest birds, most of
the mammals, and about 70% of the plants (Table 1). They are also a major source
of resins, fruits, and timber for domestic use and for export. Thus, these agroforests
are potentially a sustainable resource, a valuable compromise between conservation

Figure 1 The impact of agricultural intensification on an agroecosystem. (From Swift, M. J.
and Anderson, J. M., 1993. Biodiversity and Ecosystem Function, Schulze, E.-D.
and Mooney, H. A., Eds., Springer-Verlag, Berlin. With permission.)

Figure 2 The relationship between the two functions of trees and the three goals of agro-
forestry to meet three global challenges. (Modified from ICRAF 1997, Int’l Centre
for Research in Agroforestry Medium Term Plan 1998–2000. With permission.)
Table 1 Biodiversity in Indonesia Agroforests: Observed Numbers of Species

Primary Rubber Damar Durian Rubber
Forest Agroforest Agroforest Agroforest Plantation
Birdsa 179 105 92 69 —
Collembolab
Leaf litter 20.6 22.8 — — 11.6
Soil 13.7 16.0 — — 8.3
Mammalsc — 39 46 33 —
Treesd 171 92 — — 1
Total plantsd 382 266 — — 6
a Thiollay, 1995.
b Deharveng, 1992.
c Sibuea and Herdimansyah, 1993.
d Michon and de Foresta, 1995.

of tropical forest biodiversity and profitable use of natural resources, since in
addition to their biodiversity these multistrata damar agroforests in Sumatra are
financially attractive.
Damar resins are utilized by industries in Indonesia or exported worldwide. In
1984, the export market represented one third of the harvested volume, a trade rising
from 250 to 400 t/year between 1972 and 1983 (Michon et al., 1998). In 1994, the
damar production was expected to reach 10,000 t (Dupain, 1994), at a value of U.S.
$300 to 400/t. Of this trade, 80% is met by the damar agroforests. The economic
value of the damar trade and its associated activities is of major significance to the
villages around Krui. In 1993, the profits from damar production were U.S. $7.2
million from sales, U.S. $2.6 million from added value, and U.S. $1.4 million from
wages. To this is added U.S. $0.3 million in profits made by Krui traders (Michon
et al., 1998). This analysis excludes the locally consumed products from these
agroforests, e.g., fruits, vegetables, spices, fuelwood, timber, palm thatching, rattan,
bamboo, fibers, as well as paddy rice.
With the exception of plantation crops, many farming systems in the tropics,
including traditional subsistence swidden farming, are based on mixtures and are
frequently haphazard in their configuration and spacing. In contrast, monocultures
are particularly prevalent in countries with temperate climates. It is not clear whether
or not the tendency to complexity and random distribution of the components of
farming systems in the tropics is a deliberate attempt by farmers in the tropics to
mimic the diversity of natural ecosystems in order to minimize risk.
In contrast to traditional agroforestry, the recent development of agroforestry as
a science by agronomists and foresters has tended to adopt the temperate model and
to plant the tree component in lines, regular patterns, or along the contour of sloping
land (see review by Cooper et al., 1996). This is especially the case in countries
where farm size is large (e.g., Australia), where large areas of countryside are planted
in geometric patterns.
Modern agroforestry has also tended to be a set of stand-alone technologies, that
together form various land-use systems in which trees are sequentially or simulta-
neously integrated with crops and/or livestock (Nair, 1989). Recently, however, it
has been suggested that agroforestry practices should be successional phases in the
development of a productive and complex agroecosystem, akin to the succession of
natural ecosystems (Leakey, 1996). In this way, trees producing different products
can be used to fill niches in a mosaic of patches in the landscape, making the system
ecologically more stable and biologically more diverse. It is anticipated that this
diversity would increase with each phase of the agroecological succession.
Toward this end, current activities at ICRAF are focusing on the development
of agroforestry as “a dynamic, ecologically-based, natural resource management
system that, through the integration of trees on farms and in the landscape, diversifies
and sustains production for increased social, economic and environmental benefits.”
One aspect of this is to determine, through the use of models, the best land-use
options for agricultural productivity and biodiversity conservation: the choice
between integration or segregation (van Noordwijk et al., 1995b).
In parallel with these developments in agroforestry there has also been a move
to promote the domestication of indigenous trees, the “Cinderella” trees overlooked

by science (Leakey and Newton, 1994a; Leakey and Jaenicke, 1995; Leakey and
Izac, 1996).
Bringing the new ideas about agroforestry and about domestication together
provides one with a new paradigm for sustainable land-use development that focuses
on two aspects of biodiversity:
1. Diversifying agroecosystems
2. Capturing and enhancing intraspecific diversity
AGROFORESTRY AND THE DIVERSIFICATION OF

AGROECOSYSTEMS
From past experience, domesticated trees are frequently grown in monocultures,

but they could play an important role in species-rich multistrata agroforests (Leakey,
1996b). The development of multistrata systems that include cultivars of domesti-
cated trees could increase the profitability of these agroforests. Thus this approach
could, it seems, go a long way toward the establishment of land uses that will fulfill
the needs of rural and urban populations for food and income, while maintaining
much of the biological diversity of forests or rehabilitating degraded ecosystems.
Much research will be needed, however, to achieve this and to demonstrate that
productivity and profitability are not necessarily environmentally damaging. Evidence
already emerging from studies to develop viable alternatives to slash-and-burn agri-
culture suggests that the greenhouse gas emissions, especially methane, from areas
where sources such as paddy fields are juxtaposed with perennial vegetation are lower
than from areas monocropped with rice (van Noordwijk et al., 1995a). However, the
successful establishment of trees on cleared sites is known to suffer from changes in
the populations and species diversity of symbiotic microflora associated with land
clearance (Leakey et al., 1993; Mason and Wilson, 1994), and similar changes prob-
ably occur in the beneficial micro- and mesofauna above- and belowground. Evidence
exists for the negative effects of site clearance on soil fauna populations (Eggleton
et al., 1995) and for the need to restore them to ensure soil fertility.
A challenge for agroforestry research is to develop economically and socially
acceptable land-use systems that function like undisturbed ecosystems and maintain
biodiversity. Could complex multistrata agroforests, like those of Sumatra, be devel-
oped in humid West Africa and in Latin America? The answer is almost certainly
yes. Indeed, simple indigenous multistrata systems already exist, such as the cocoa
farm, and the compound gardens of West Africa (Okafor and Fernandes, 1987),
while in the Peruvian Amazon, multistrata agroforests have been found to be an
economically attractive system (Table 2).
DEVELOPMENT OF MULTISTRATA AGROFORESTS
There are plenty of tree species that have traditionally provided local people
with their daily needs for the full range of nontimber forest products, which could

Table 2 Comparison of Net Present Values and Internal Rates of Returns of
Production Systems in 1985 and 1991 Prices Using a 15-Year Time
Horizon in Yurimaguas, Peru
1985 Prices 1991 Prices
Net Present Internal Net Present Internal
Value Rate of Value Rate of
Production Option (U.S.$ ha–1) Return (%) (U.S.$ ha–1) Return (%)
Multistrata system 6444 219 6727 831
Peach palm 979 35 2061 64
Shifting cultivation 79 7 218 19
Note: Low input and high input continuous cultivation both had negative net
present values.
From ICRAF, 1994 Annual Report, Nairobi, Kenya, 1995.
now be incorporated into agroforestry systems. Table 3 illustrates candidate species

for West Africa (see also Okafor and Lamb, 1994; Abbiw, 1990). How can these
complex agroforests be further developed in the tropics? Probably three things are
required. First, there is the need to identify the species of commercial importance
of relevance to local markets. Second, there is a need to develop an entrepreneurial
mentality among the community, who have traditionally been subsistence farmers
and hunter–gatherers. Third, there is a need to determine how best the tree species
may be combined to develop agroforests. The growth of the urban market and the
absence of jobs in the urban areas may provide the commercial incentive required.
What is probably missing is the demonstration of what is possible, particularly in
the areas near urban markets.
With such a wide choice of species for the middle and upper strata of multistrata
agroforests, clearly research to determine the best combinations and configurations
is much needed, but also extremely difficult. There are, therefore, three research
approaches that can be taken:
1. The testing of prototype systems (i.e., best-guess combinations), perhaps aimed at

market needs, and developed with the help of farmers with some experience of
compound gardens;
2. Research to test specific hypotheses aimed at the development of some principles
regarding the optimal combinations and/or densities of trees in the different strata,
which involves both complementarity of species biologically and in terms of labor
demands and market opportunities;
3. Use of random mixtures of the species in unstructured combinations, as would
probably be developed by farmers.
Research is needed to determine whether or not the apparently random distribution

of trees in many existing examples of multistrata systems is indeed random, or
whether farmers from experience grow species in certain combinations. Understand-
ing this process would be of benefit in assisting attempts to know why multistrata
systems evolve differently under different social and ecological conditions and would
therefore help to transfer these systems more effectively to new areas.

Table 3 A Sample of the West African Tree/Shrub/Liane Species Appropriate for
Growth in Multistrata Agroforests and for Domestication
Mature Height
Common Names (m)
Anthocleista schweinfurthii Ayinda 15–20
Antrocaryon micraster Aprokuma/onzabili 40–50
Baillonella toxisperma Moabi 45–55
Calamus spp. Rattan 35–45
Canarium schweinfurthii Aiele/Africa canarium/incense tree 45–55
Chrysophyllum albidum Star apple 30–40
Cola acuminata Kola nut 15–25
Cola lepidota Monkey kola 10–20
Cola nitida Kola nut 20–30
Coula edulis Coula nut/African walnut 25–35
Dacryodes edulis African plum/Safoutier 15–25
Entandrophragma spp. Sapele/tiama/utile/sipo 50–60
Garcinia kola Bitter kola 20–30
Gnetum africanum Ero 0–10
Irvingia gabonensis Bush mango/andok 20–30
Khaya spp. African mahogany 50–60
Lovoa trichiloides Bibolo/African walnut 40–50
Milicia excelsa Iroko/mvule/odum 45–55
Nauclea diderichii Opepe/kusia/bilinga 35–45
Pentaclethra macrophylla Oil bean tree/Mubala/Ebé 20–30
Raphia hookeri and other spp. Raphia palm 5–15
Ricinodendron heudelotii Groundnut tree/nyangsang/essessang 40–50
Terminalia ivorensis Framiré/Idigbo 45–55
Terminalia superba Fraké/afara/limba 45–55
Tetrapleura tetraptera Prekese/Akpa 20–30
Treculia africana African breadfruit/etoup 20–30
Trichoscypha arborea Anaku 15–25
Triplochiton scleroxylon Ayous/obeche/wawa 55–65
Vernonia amydalina Bitter leaf 0–10
Xylopia aethiopica Spice tree 15–25
From Leakey, R. R. B., Agroforestry Systems, 1998. With permission.
There are a number of options on how to apply these approaches to the devel-
opment of multistrata systems:
1. Enrichment planting within logged or degraded forest;

2. Planting in cleared forest land as a perennial tree alternative to slash-and-burn
agriculture;
3. Planting under a plantation of either an upper or middle strata species.
Of these options, 1 and 3 have merit environmentally in that the system will be
developed more quickly as a demonstration and will not leave the land bare at the
establishment phase. On the other hand, option 2, the currently most practical from
the perspective of smallholders, is probably the most relevant in terms of developing

multistrata systems. Such systems are both relevant on good soils of land cleared at
the forest margin and on already degraded land, perhaps with proximity to urban
markets. Because this option is the most relevant practically, it is also probably the
one on which the greatest research effort should be concentrated.
In all these options, research is especially required to determine the functional
groups of species which will do well in the lower strata, where light will be a limiting
factor. Currently in Southeast Asia, staple food crops are grown beside the multistrata
agroforests, because they are light demanding. This may be the best arrangement,
but it is also possible that new crops, or new cultivars of existing crops, could be
integrated into multistrata agroforests.
BIODIVERSE AGROECOSYSTEMS
From the biodiversity viewpoint, there is a difference in agroecosystems between

the planned biodiversity and the unplanned, or associated, biodiversity. The latter
are all those organisms, above- and belowground, that have found niches to fill
among the planted trees and crops. The extent to which unplanned biodiversity
occurs in different agroecosystems is not well known or understood, although studies
have started to address the effects of a broad spectrum of agricultural practices on
wildlife populations (McLaughlin and Mineau, 1995; Perfecto and Snelling, 1995).
Swift et al. (1996) have, however, drawn four very different scenarios for the
relationships between agricultural intensification and biodiversity (Figure 3),
although these may be very scale dependent (e.g., from farm to landscape) and
probably also vary depending on the level of biodiversity at the time of planting.
Thus, the biodiversity associated with an agroforest planted on recently cleared land
at the forest margin, as an alternative to slash-and-burn agriculture, would almost
certainly be very different from the same tree/crop mixture planted to rehabilitate
already degraded land. There is a need for controlled experiments to determine these
relationships between intensification and biodiversity. In addition there is a need to
determine the patterns of diversity in different agroforestry systems and their impli-
cations for ecological functioning at different scales. There is currently unresolved
debate about the functional role of species diversity in ecosystems (Johnson et al.,
1996), which makes it difficult to suggest best practices. However, it seems that
planned biodiversity should aim at maximizing ecosystem processes (nutrient
cycling, production of different products, light requirements, etc.) and structural
complexity, rather than increasing the number of species per se. However, species
numbers may also impact on function. The few studies in which the diversity of
agricultural ecosystems has been manipulated suggest that increases in diversity
from 0 to 10 plant species alters ecosystem function, but that there is little effect
beyond that point (Schulze and Mooney, 1993).
Species also vary in their importance in different food webs, where, once again,
scale is important. For example, for a fruit tree species to support a population of
monkeys will require a large habitat. A small area of fruit trees may therefore avoid
monkey damage and so be preferable for production, but without a large area there
may not be a market for the crop. Thus, ecological and economic factors affecting

Figure 3 Hypothetical relationships between agricultural intensification and agroecosystem
biodiversity. (From Swift, M. J. et al., 1996. Functional Roles of Biodiversity: A Global
Perspective, Mooney, H. A. et al., Eds., SCOPE 55, John Wiley & Sons, New York.
With permission.)
the viability of production systems can be antagonistic. Typically, predators require

larger range than their prey; thus for the sustainable production of these fruits the
area may need to be big enough to support leopards. Top predators are, however,
often unwelcome in farmland while their absence creates pest management issues
for farmers. In this context, a land-use mosaic with corridors between forest patches
may be the most appropriate means of acquiring the scale needed to achieve some
level of ecological equilibrium; however, the likely benefits of corridors between
forest patches are not fully understood (Hobbs, 1992). This concept thus returns to
issues of scale, land-use design, and the “integrate vs. segregate” debate (van Noord-
wijk et al., 1995b) in developing optimal land-use strategies.
FOREST PATCHES, BIOGEOGRAPHICAL ISLANDS,

AND AGROFORESTRY
In the tropics, conservationists have focused their attention on the protection of

natural forests and woodlands and, until recently (Schelas and Greenberg, 1996),
have not given much attention to the widely dispersed forest patches throughout
human-occupied landscapes. These patches are often critical components of a
farmer’s environment, being a source of products and environmental services of
importance to the farmer’s livelihood and welfare. As biogeographical islands, their
role in maintaining biological diversity is also crucial. Issues of scale are central to
this role, and thus, in a landscape mosaic, forest patches and areas of agroforestry
are potentially complementary, especially when considering the need for ecological

equilibrium and population size vis à vis genetic diversity. There is thus a need for
ecologists and geneticists to become more involved in agroforestry research and the
process of farmer reforestation. Biological and genetic diversity are, for example,
becoming an issue in the densely populated, high-potential areas of Kenya. These
are areas where the situation of “more people, less erosion” (Tiffen et al., 1994) is
also becoming “more people, more trees” (Holmgren et al., 1994), as tree biomass
is increasing at 4.7%/year, while the high human population is expanding at 3.0%.
However, of concern is the fact that about half the trees planted on a farm are of a
single species (Grevillia robusta), while the other half consists of about 10 to 20
species. This raises the question at what point the tree population reaches a size and
distribution such that the controls that prevent isolated outbreaks of pests and disease
break down. Not enough is known about the patch dynamics of such outbreaks
relative to species numbers and configurations to steer agroforesters in the develop-
ment of risk-averse strategies. The use of only a few tree species in agroforestry to
provide corridors between forest patches could obviously allow their pests and
diseases the opportunity to spread, while reducing the effectiveness of corridors for
the ecological and genetic stability of other species. This reaffirms the need for both
diversity and the need to consider the ecological implications of landscape design.
In the case of G. robusta, which was introduced into East Africa from Australia, the
risks of its dominance in the landscape could be exacerbated by its limited genetic
base. Fortunately, new and more diverse accessions of G. robusta have now been
introduced in East Africa, but it will take some time for these to be disseminated
around the countryside. This example illustrates the need to ensure that agroforestry
promotes the importance of diversity and not just the planting of trees by farmers.
Strictly, biogeographical islands are completely isolated populations, in which
island size is positively related to biodiversity. The concept has, however, also been
applied to shrinking habitats such as woodlands in farmland. Although Simberloff
(1988) concluded that, in a wildlife conservation context, treating woodlands as
islands in this way provided contradictory or inconclusive evidence, there does
seem to be some merit in considering (1) how agroforestry could perhaps benefit
biodiversity through increasing island sizes and (2) what might be the possible
impacts of such action. It is not known, however, if the species–area relationships
apply when habitats are being expanded. Can agroforestry, as a form of restoration
ecology, reverse the 18 factors (e.g., low population density, inbreeding, catastro-
phe, competition, habitat destruction, disease, etc.) associated with species extinc-
tion listed by Soulé (1983)? Agroforestry can probably reverse some of them, but
which of the 18 factors are the most crucial and which should these be the prime
targets for the development of a healthy agroecosystem? At present, it is only
genetic diversity that is getting any attention. The decrease in size of biogeograph-
ical islands lowers genetic diversity, increases the likelihood of inbreeding, and
lowers the chances of future allopatric speciation (Soulé, 1986). Agroforestry,
because of its production orientation, is promoting the retention and introduction
of genetic diversity among the tree species planted by farmers. This aspect of tree
domestication at least is one positive step toward a healthier and more productive
environment. It is hoped that, as agroforestry is increasingly seen as applied ecology,

other such steps will follow, such as the geographic expansion of the multistrata
agroforestry systems already described.
DOMESTICATION OF TREES FOR TIMBER AND

NONTIMBER FOREST PRODUCTS
Throughout the tropics there are numerous perennial woody species that have
provided indigenous peoples with many of their daily needs for millennia. Many
of these people have now left the land for urban life, but they still demand traditional
food, medicines, and other natural products. These traditionally important woody
plants are virtually undomesticated. These neglected “Cinderella” species have
great genetic diversity and also play a key role in biological, chemical, and hydro-
logical cycles, protecting soils and providing ecological niches. The food-produc-
ing species are also important for food security, especially in the dry season, as
well as a source of vitamins and minerals critical for the health and nutrition of
children and pregnant women.
There are four groups of wild trees, shrubs, and lianes which could be rapidly
domesticated for agroforestry and which can be viewed as potentially important
sources of income for farmers. These trees produce
1. The traditionally important wild foods, mostly fruits, nuts and leaves for vegetables;
2. The traditionally important fibers;
3. Locally and industrially important pharmaceuticals and other extractives such as
gums and resins;
4. Commercially important quality timbers and woods.
The domestication of tree species is a dynamic process which develops from

deciding which species to domesticate and proceeds through background socioeco-
nomic studies, the collection of germplasm, genetic selection and improvement to
the integration of domesticated species in land-use systems (Leakey and Newton,
1994b; Leakey and Jaenicke, 1995). Domestication is an ongoing process in which
genetic and cultivation improvements are continuously refined. In genetic terms,
domestication is accelerated and human-induced evolution. Domestication, however,
is not only about selection. Simons et al. (1998) contend that it integrates the four
key processes of the identification, production, management, and adoption of tree
genetic resources.
Strategies for tree domestication will vary depending on the value of the products,
the extent of intraspecific variation, and many other factors, but for high-value
species such as those producing marketable forest products the vegetative propaga-
tion of superior genotypes identified from within the existing wild populations will
be appropriate (Leakey and Simons, 1998a). This is the approach generally followed
in horticulture. Thus, an individual with superior yield, fruit flavor, stem form, or
wood quality can be mass produced by vegetative propagation. In this way it is
possible to select those clones likely to develop above-average characteristics in any

given trait. By a series of ongoing selections and an ever-increasing intensity of
selection, it is also possible to achieve rapid and substantial genetic improvements.
In situations where desirable traits are not easily identified, it is possible to multiply
a number of copies of each plant and to establish these in field trials and then to
observe the development of distinct genetic differences between each clone (Long-
man and Jeník, 1987).
When identifying and selecting trees for cloning by vegetative propagation,
there are two important criteria. It is important, first, to ensure that those that are
highly superior for the desired traits are chosen and, second, to ensure that the
selected individuals are unrelated and as genetically diverse in other traits as
possible. This calls for a risk-aversion strategy (see Leakey, 1991) that allows
intensive selection (e.g., 1 out of 100 to 10,000 trees) from among trees from
different populations, ideally from throughout the range of the species. Furthermore,
this should be an ongoing rolling program of multiple-trait selection, in which more
and more intensive selection is imposed through the addition of selection criteria
for new traits while, at the same time, new sources of genetic stock are continually
added as new accessions enter the program. These new accessions should come
from further germ plasm exploration and from breeding programs and so continually
broaden the genetic base of the planted trees. Molecular genetics techniques can
be used to ensure that genetic diversity is maintained in the clones used for
commercial production.
Selection procedures will vary depending on the product and the situation. For
example, for indigenous fruits, rapid progress will be made if indigenous knowledge
can be used. Usually rural people know which are the best individual trees in their
area for yield, fruit size, or flavor. Thus, as with temperate apples, pears, etc., people
can be asked to report the existence of superior trees, so reducing the task of
screening large numbers of trees. On the other hand, for medicinal trees, it is more
likely that a chemical screening process will be required, but the magnitude of this
task can probably be reduced by starting on a population basis, since it is likely that
trees from certain environments will be richer in the required metabolites. Mean-
while, for timber trees, log size and straightness are the first selection criteria. Various
forms of “plus-tree” (i.e., elite genotype) provenance and progeny selection are well
known. To these have recently been added some “predictive tests” which can be
applied in the nursery as a procedure for mass screening from genetically diverse
seedling populations.
COMMERCIALIZATION
For the success of a domestication program, there must be markets for the
products. Many of the products from tropical trees are already sold on local and
regional markets, and a few have broken through into the international marketplace.
For example, while the pulp of the bush mango (Irvingia gabonensis) is eaten fresh,
its extracted kernels and those of the related species, I. wombolu, are traded region-
ally throughout the year. From Cameroon, this trade extends to Nigeria, Equatorial
Guinea, and Gabon (Falconer, 1990; Ndoye, 1995). Similarly, the trade of kola nuts

extends from humid zone countries of West Africa up into the dry zone where there
is a big demand by the Muslim community (Falconer, 1990). Chewing sticks
likewise are traded northwards in West Africa, with a street value put on the trade
from Kumasi market in Ghana of about U.S. $9 million/year (Falconer, 1992).
Again, in Cameroon the bark of Prunus africana (Pygeum) is exported to Europe
where pharmaceutical products estimated to be worth $150 million/year are pro-
duced for worldwide trade and treatment of prostate gland disorders (Cunningham
and Mbenkum, 1993). In southern Africa, some indigenous fruits are marketed
locally as wines and jams, with a liqueur from Sclerocarya birrea fruits (Amarula)
now on the international market. In Amazonia, products from the peach palm
(Bactris gasipaes) are also being exported. The palm heart trade has been estimated
at around $50 million/year (Clement and Villachica, 1994), with the fruits also
having a similar value.
Commercialization is both necessary and potentially harmful to farmers. Without
it the market for products is small. On the other hand, it is potentially harmful to
rural people if it expands to the point that outsiders with capital to invest come in
and develop large-scale monocultural plantations for export markets.
Leakey and Izac (1996) have examined some of the issues requiring consideration
to ensure the economic viability of small-scale production and under what conditions
small-scale production can be competitive with large-scale production.
TREE DOMESTICATION IN PROGRESS
Since 1993, ICRAF has initiated programs to identify priority trees for domes-
tication (Franzel et al., 1996) and started germplasm collection in the humid lowlands
of West Africa, the semiarid lowlands of West Africa, the Miombo Woodlands of
the southern Africa plateau, and in western Amazonia (Leakey and Simons, 1998b).
Through these programs ICRAF is developing the concepts, strategies, and policies
associated with capturing genetic diversity of a wide range of indigenous trees for
growth in agroforestry systems. These will be developed as the programs evolve. In
this way, it is hoped the promises of agroforestry to alleviate poverty and to mitigate
environmental degradation and the loss of biodiversity will to be fulfilled.
CONCLUSIONS
The vision of agroforestry presented here is as an integrated land use that, through
the capture of intraspecific diversity and the diversification of species on the farm,
combines increases in productivity and income generation with environmental reha-
bilitation and the creation of biodiverse agroecosystems. In most places this is just
a vision, but there are increasing numbers of examples where the vision is already
a reality. The body of ecological data from agroforestry research is growing, and
the research agenda is changing toward systems thinking. There is much to do to
encourage these developments and the socioeconomic/policy conditions that promote
them. There is, however, an urgent need for the reasons for hope to be heard above

the shouts of doom and gloom about tropical forests, if the needs of growing human
populations are to be met without sacrificing the biological diversity that keeps our
ecosystems functional.
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CHAPTER 9
The Role of Agroecosystems in

Wildlife Biodiversity
Thomas E. Lacher, Jr., R. Douglas Slack, Lara M. Coburn, and

Michael I. Goldstein
CONTENTS
Introduction: The Interaction Between Wildlife and Agroecosystems

Effects of Agroecosystems on Wildlife
Positive Effects of Agriculture on Wildlife
Negative Effects of Agriculture on Wildlife
Case Studies: The Use of Agroecosystems by Wildlife
Wildlife and Rice Cultivation
Migratory Birds, Agroecosystems, and Agricultural Chemicals
Agricultural Practices in Coffee Agroecosystems
Trees as Row Crops: Plantation Forestry and Wildlife
Conclusions — Net Effects of Agroecosystems
Loss of Biodiversity
Change in Community Structure
Recommendations for the Mitigation of Impacts
References
INTRODUCTION: THE INTERACTION BETWEEN WILDLIFE

AND AGROECOSYSTEMS
Agriculture is among the most important of all human enterprises. A small

number of species of crops, domesticated by a variety of early civilizations, now

provides the basis of most of our food consumption. Fifteen species of plants,
primarily grains, provide over 90% of all human energy needs, and over 98% of all
human food is produced in terrestrial habitats (Paoletti et al., 1992). Agriculture,
forestry, and human settlements occupy 95% of all terrestrial environments, whereas
nondeveloped areas such as national parks account for only 3.2% worldwide (Pimen-
tal et al., 1992). The balance between protected areas and modified landscapes has
shifted strongly toward the latter, and there is little doubt that agricultural diversifi-
cation and expansion has decreased biodiversity over the past two centuries (Dahl-
berg, 1992). Thus, concerns over the effects of agriculture on wildlife have increased
in recent years.
The major impacts on wildlife are caused by habitat conversion and habitat
fragmentation. The U.S. provides a good example of this process in a developed
country. About 70% of the U.S. (excluding Alaska) is held in private ownership by
millions of individuals, although 50% of the land is in the hands of only 2% of the
population. About 50% of the country is either cropland, pasture land, or rangeland,
owned by approximately 4.7 million individuals (U.S. Department of Agriculture,
1996). Over 200 different species of crops are produced on this land; however, 80%
of this total production is accounted for by four species: hay, wheat, corn, and
soybeans (U.S. Department of Agriculture, 1996). When forests or grasslands are
converted to an agroecosystem, virtually all native species of plants and many of
the animals are lost. There is often some degree of utilization of the agricultural
fields by vertebrates and invertebrates, but when these species cause losses of crops,
they are controlled, usually by chemical means. This often results in the elimination
of nontarget organisms as well. Some natural ecosystems, for example, wetlands,
have been particularly severely impacted by agricultural expansion. Up until the
1950s, approximately 87% of all wetland conversion was attributable to agriculture,
though recent legislation has reduced that percentage. In fact, between 1982 and
1992, 57% of wetland losses were due to urban expansion, and only 20% to agri-
culture (U.S. Department of Agriculture, 1996).
Economic incentives frequently contribute to habitat conversion. Increased eco-
nomic pressures and new technological innovations can cause losses of biological
diversity in the early stages of development (Howitt, 1995). This is especially a
problem in the tropics. Developed country policies often determine the agricultural
practices of developing countries; developing countries generally function as “price
takers” and are largely exporters of primary products to the developed world
(McNeely and Norgaard, 1992). Agricultural development projects financed through
international aid agencies have neglected environmental issues in the past. The
impact on wildlife in developing tropical nations has been substantial; however, there
are several innovative proposals to link economic and ecological systems in agri-
cultural development (McNeely and Norgaard, 1992).
We present a summary of the effects of agriculture on wildlife, both positive
and negative. For each individual scenario, there are both benefits and costs. We
present four different case studies that attempt to capture the complexity of issues
and effects. Finally, we close with some recommendations.

EFFECTS OF AGROECOSYSTEMS ON WILDLIFE
Positive Effects of Agriculture on Wildlife
Several aspects of agroecosystems can positively affect wildlife populations. One

aspect of the fragmentation of an agroecosystems/forest mosaic is the creation of
edge habitat. This results in the edge effect, or the tendency for the variety and density
of some species of plants and animals to increase at the border between different
plant communities (Forman, 1997). Edges, or ecotones, contain species from both
habitats as well as a subset of species considered to be edge specialists (Yahner,
1988). Some of these species are important game species (e.g., white-tailed deer in
North America), which has resulted in a management practice among game biologists
of creating edge habitat (Yoakum and Dasmann, 1971). The cost of edge creation is
a reduction in the amount of forested habitat available and a decline in the abundance
and richness of forest species if fragmentation becomes too severe. There are, how-
ever, some species that exist and even thrive in altered or fragmented habitats,
especially those that have small area requirements or that are mobile and can easily
move among habitat patches (Merriam, 1991; Noss and Cooperrider, 1994).
Grain agriculture often leaves residual seeds on the ground after harvest that serve
as a valuable resource for many species of wildlife. Rice plantations provide rice
grains as food as well as surrogate wetlands for many species of waterfowl (see below
for a detailed case study). In the U.S., concern over the environmental impact of
agriculture has also led to the passage of legislation geared toward the enhancement
of wildlife habitat. Although not specifically tied to agriculture, the Endangered
Species Act of 1973 has led to changes in agricultural practices when species of
concern were potentially impacted. For example, concerns over the impact of irriga-
tion on salmon fisheries led to the restriction of access to federally supplied irrigation
water in California (Day, 1996). The Farm Act of 1996 created several valuable
programs for the protection of wildlife (Table 1) and has included the conservation
Table 1 Programs and Provisions of the 1996 Federal

Agriculture Improvement Act (Farm Act)
1. Environmental Quality Incentives Program
2. Wetlands Reserve Program and Conservation Reserve Program
3. Farmland Protection Program
4. Swampbuster and Wetland Provisions
5. Wildlife Habitat Incentives Program
6. Flood Risk Reduction Program
7. Emergency Watershed Protection Program
8. Conservation of Private Grazing Land
9. National Natural Resources Conservation Foundation
10. Conservation Farm Option
11. State Technical Committees
Source: U.S. Department of Agriculture, America’s Private Land: A
Geography of Hope, U.S. Department of Agriculture, Washington, D.C.,
1996.

of wildlife habitat as a goal of agricultural programs. For example, the Wildlife
Incentives Program was the first agricultural program developed exclusively for the
creation and protection of wildlife habitat. The Conservation Reserve Program was
originally presented in the 1985 Farm Act and reauthorized, in 1996, the setting aside
or converting of as much as 36.4 million acres of environmentally sensitive farmland,
through 2001. The Wildlife Habitat Incentives Program of 1996 set aside $200 million
for restoration programs in the Everglades Agricultural Area provision (Day, 1996).
All of this legislation, explicitly linked to agriculture, has benefited wildlife.
Negative Effects of Agriculture on Wildlife
The greatest negative effect of agriculture on wildlife is the conversion of natural

vegetation to an agroecosystem. Habitat loss directly reduces biodiversity. At least
71 species and subspecies of vertebrates1 and at least 217 species of plants2 have
gone extinct in North America (north of Mexico) since the arrival of Europeans.
Over 95% of our original virgin forests are now gone from the lower 48 states (Postel
and Ryan, 1991). The decline in large mammalian predators as a result of habitat
loss to agriculture has resulted in an increase in the population of deer, which have
now become an agricultural pest in many regions (Day, 1996). These losses of species
and habitats are the result of the gamut of human activities, of which agricultural
activities form a major part.
Given that a certain amount of land will be dedicated to providing for human
food and nutrition, the next most significant effect is fragmentation. Habitat frag-
mentation is defined as the subdivision of continuous habitat over time; the most
important large-scale cause of habitat fragmentation is expansion and intensification
of human land use (Burgess and Sharpe, 1981; Harris, 1984). Fragmentation is
considered to be an important cause of local extinction (Wilcox and Murphy, 1986).
Fragmentation results in a loss of original habitat, a reduction in the sizes of the
patches of remaining habitat, and an increase in the degree of interpatch distances,
all of which increase the rate of local extinction (Harris, 1984; Wilcove et al., 1986).
Even certain aspects of the edge effect, positive for some species, are detrimental
to others. Edges can serve as potential ecological traps for breeding birds by con-
centrating nests in a small area where the risk of predation is high (Rudnicky and
Hunter, 1993). There is also an apparently high rate of nest parasitism of breeding
birds by cowbirds in edge habitats (Brittingham and Temple, 1983).
Activities other than deforestation associated with agriculture also pose threats
to wildlife. The use of agricultural chemicals expanded after the end of World War
II and their impact on avian populations became a national issue after the publication
of Rachel Carson’s Silent Spring in 1962. Several recent review volumes have
addressed the impacts of agricultural chemicals on wildlife and discussed the
attempts to mitigate these impacts (Kendall and Lacher, 1994; Colborn et al., 1996).
1 See The Nature Conservancy, 1992. Extinct Vertebrate Species in North America, unpublished draft list,
March 4, 1992. The Nature Conservancy, Arlington, VA.

2 See Russell, C. and Morse, L., 1992. Extinct and Possibly Extinct Plant Species of the United States
and Canada, unpublished report, review draft, 13 March 1992, The Nature Conservancy, Arlington, VA.

Although persistent pesticides, like DDT, which was an issue in Carson’s time, are
rarely used, other more acutely toxic compounds now pose a threat of mortality to
wildlife. Others, so-called endocrine disrupters, may cause long-term reproductive
damage (Colborn et al., 1996). Some of the most significant victims of pesticides
have been nontarget species of insects. The loss of these potential pollinators will
have far-reaching effects, even on many agricultural crops (Buchmann and Nabhan,
1996). States like California, which was once the largest U.S. user of agricultural
toxicants, now have ambitious programs to reduce chemical use (Anderson, 1995).
This includes compounds recently suspected of acting as endocrine-disrupting chem-
icals (Fry, 1995).
Noss and Cooperrider (1994) present a series of summary tables in chapter 3 of
their book on the impacts of a variety of land-use practices. Concern for increased
rates of local extinction and the concomitant loss of biodiversity as a result of
agricultural development is a growing issue in tropical regions as well as temperate
zones (Holloway, 1991).
CASE STUDIES: THE USE OF AGROECOSYSTEMS BY WILDLIFE
Wildlife and Rice Cultivation
In the U.S., Texas is one of seven states that grow rice (Texas Rice Task Force,
1993). The Texas rice crop is grown in the gulf prairies and marshes of the upper
Texas coast (Gould, 1975). The native tall grass prairies historically extended inland
from extensive coastal marshes for approximately 20 to 150 km. The prairies were
characterized by nearly level to gently sloping topography interspersed with small,
rain-filled depressions. Prior to the 1900s, the prairies of the upper Texas coast were
grazed by herds of bison (Bison bison) and wild horses (Robertson and Slack, 1995).
As the land was settled, bison and wild horses were replaced with free-ranging cattle
and later with agricultural crops (Craigmiles, 1975; Stutzenbaker and Weller, 1989).
Rice was first introduced to the coastal prairies in the mid-1800s. By 1954, a peak
of 254,000 ha of rice were harvested on the gulf prairies (Hobaugh et al., 1989).
Currently, about 110,000 ha of rice are harvested in Texas producing an aggregate
addition to the Texas economy of almost $1 billion (Texas Rice Task Force, 1993).
In addition, the economy of the rice growing region of the state is enhanced by
significant expenditures for recreational hunting.
Rice fields are prepared for planting in late winter with actual planting occurring
in March or April. Fields are flooded shortly thereafter until immediately prior to
harvest in August. These flooded fields provided large expanses of wetlands for some
resident birds to use. In portions of the Texas rice belt a second crop (“ratoon crop”)
results from resprouting from the initial planting and is harvested in October. Har-
vested fields contain waste grain and are left to stand fallow for up to 2 years. During
the subsequent seasons, the fallow fields are grazed by cattle. Therefore, a typical,
3-year, rice–pasture rotation system involves three fields; during early winter rice is

harvested in one field, another field is plowed in preparation for planting rice the
next spring, and the third field is being grazed (Hobaugh et al., 1989).
At a landscape scale, the tall grass prairies of the upper Texas coast were a
relatively homogeneous matrix of tall prairie grasses with small, scattered, natural
depressions (Figure 1a). At smaller scales within the matrix, the landscape was
heterogeneous, with grasses, forbs, and scattered natural wetlands with associated
aquatic vegetation. Because of the intensive rice-cropping system, the resulting
landscape is a reversed image of the native tall grass prairie environment (Figure
1b) — a heterogeneous mosaic at the landscape scale, with homogenous field-sized
stands of vegetation, prepared fields, or pastures.
The rice-cropping system in Texas lies adjacent to a heavily industrialized region
with over 30% of the U.S. petroleum industry and more than 50% of the U.S.
chemical production occurring in this region (Robertson and Slack, 1995). In addi-
tion, the Houston–Galveston metropolitan area is the fourth largest metropolitan area
in the U.S. As a result of these economic pressures, the area of wetland habitats has
declined dramatically in Texas, and especially in coastal regions of the state (Ander-
son, 1996; Moulton et al., 1997). Current estimates of losses show that >35% (84,000
ha) of Texas’ coastal marshes have been destroyed since the 1950s (Anderson, 1996).
The net effect of the landscape mosaic produced by the Texas rice-cropping
system has been a dramatic change in use by migratory birds since the advent of
rice agriculture. Lesser snow geese (Chen caerulescen), greater white-fronted geese
(Anser albifrons), and Canada geese (Branta canadensis) only began to use the
prairie after rice agriculture became established on the upper Texas coast (Hobaugh
et al., 1989). Waterfowl were commonly associated with the small natural depres-
sions in the native prairies (McIlhenny, 1932). However, it wasn’t until the 1940s
and 1950s with mechanization of rice farming, extensive irrigation, and the 3-year
rice rotation system that geese and waterfowl began to exploit the system fully
(Hobaugh et al., 1989; Robertson and Slack, 1995). Waterfowl and geese are attracted
to the mosaic of habitats because of the availability of waste grain after harvest in
the fall and the extensive areas of standing and impounded water associated with
roost ponds. Gawlik (1994) documented as much as 87 kg/ha in stubble fields
immediately after harvest. Waste rice is an important source of food for wintering
ducks, geese, and numerous granivorous passerines (Terry, 1996). Similarly, the
importance of waste rice to wintering waterfowl has been documented for the Central
Valley of California (Alisauskas et al., 1988; Brouder and Hill, 1995; Gawlik, 1994).
In addition to rice grains, green vegetation emerging during the winter in harvested
rice fields and in fields that had been prepared for the rice crop the following season,
becomes an important source of food for geese (Hobaugh, 1985; Gawlik, 1994).
Well over 2 million waterfowl and geese winter on the upper Texas coast with the
bulk of these birds found using freshwater wetlands associated with rice agriculture
(Haskins, 1996). The extensive use of rice-cultivated land by wintering lesser snow
geese has been identified as a significant component of the observed high population
growth rates. These high population densities have resulted in significant alteration
to Arctic coastal salt marsh plant communities (Abraham and Jefferies, 1997).

Figure 1 Schematic representation of the Texas coastal tall grass prairie (A) before the advent
of rice cultivation (mid-1980s) and (B) after rice production. Rice production has
fragmented the larger landscape into patches of homogeneous stands of vegetation,
prepared fields, or pastures but has greatly reduced landscape heterogeneity at
small spatial scales.
McFarlane (1994) and Terry (1996) have documented the use of the rice system
by more than 70 species of birds during an annual cycle. Most species were asso-
ciated with wetland habitats such as roost ponds, flooded rice fields, and natural
depressions. Sheridan et al. (1989) have documented at least 22 species of colonial-
nesting waterbirds nesting in 42 colonies located on the upper Texas coast including
rice lands. In addition, over 35 species of migratory shorebirds were documented
for the upper Texas coast, with 16 species found by Terry (1996). Migratory shore
birds take advantage of the wetland habitats associated with rice agriculture, as well
as moist, open fields prepared for next year’s rice crop.

Migratory Birds, Agroecosystems, and Agricultural Chemicals
Swainson’s hawks (Buteo swainsoni) are long-distance migrants whose habits

in North America have been well documented (England et al., 1997). Breeding
habitat in North America consists of open grassland and shrub steppe semiarid
ecosystems from Mexico to the prairie provinces of Canada. Birds nest in trees
adjacent to large fields, often utilizing agricultural grassland habitat for locating food
and other daily activities (Bloom, 1980). Hawks hunt on the ground or in midair,
opportunistically eating insects, small mammals, reptiles, and birds (Bednarz, 1986).
Adults primarily feed on mammals and birds to supply the nutritional requirements
to growing nestlings. Fledglings, aggregates of nonbreeding hawks, and aggregates
of premigratory mixed-age hawks forage primarily for insects such as grasshoppers
and dragonflies (Woffinden, 1986; Johnson et al., 1987).
Breeding in western North America during the boreal summer, Swainson’s
hawks migrate to the nonbreeding grounds in South America with the advancing
austral summer, maintaining both climatic and habitat similarities (Figure 2). The
journey of up to 10,000 km in each direction takes less than 2 months, and, once
settled in southern South America, hawks generally reside in the agricultural grass-
lands of the Argentine pampas (White et al., 1989; Woodbridge et al., 1995; Gold-
stein, 1997). This habitat is similar to agricultural prairies found throughout their
North American range.
Utilizing agricultural areas west and north of the capital city of Buenos Aires
during the nonbreeding season, Swainson’s hawks were found in the Argentine
provinces of La Pampa, Cordoba, Buenos Aires, Santa Fe, and San Luis (Goldstein,
1997). Hawks were encountered sunbathing and foraging in freshly tilled fields or
in fields whose crop height was less than 40 cm. Crops used included alfalfa, corn,
sorghum, soybean, and sunflower. Flocks of hawks followed insect outbreaks, trav-
eling over a small region as stages in crop growth changed throughout the season.
Other insectivorous species, such as the Chimango caracara (Milvago chimango),
burrowing owl (Athene cunicularia), Franklin’s gull (Larus pipixcan), and southern
lapwing (Vanellus chilensis) followed these insect outbreaks as well. Aplomado
falcons (Falco femoralis) also feed on agricultural sites in the pampas, predating
insectivorous songbirds.
Increasing monoculture and more intensively managing alfalfa in these regions
have also resulted in heavy reliance on agrochemicals for crop protection from insect
pests. Subsequent hot and dry conditions of the pampas during the austral summers
of 1994–95 and 1995–96 led to severe grasshopper outbreaks, exacerbating the
problem of reliance on chemical controls. Typically, the inexpensive organophos-
phate insecticide monocrotophos (MCP) was used for grasshopper controls. During
this time, when insect outbreaks and chemical controls were at their maximum, the
largest flocks of Swainson’s hawks, up to 12,000 birds, were seen.
Agrochemical controls during the austral summers from 1994 through 1996 led
to 19 documented hawk mortality incidents, accounting for approximately 6000 dead
Swainson’s hawks over two seasons (Goldstein et al., 1996; Goldstein, 1997). Hawks
died in fields while foraging for grasshoppers, in roosts after returning from foraging
bouts, and along the trajectory from fields to roosting trees. The agrochemical MCP

Figure 2 Swainson’s hawk (Buteo swainsoni) breeding range, migratory route, and nonbreed-
ing range, with a list of the common names for the Swainson’s hawk used across
the Americas. (Courtesy of M. Fuller, unpublished data.)
was determined responsible for mortality in birds from all 6 sites sampled and in
17 of 19 sites overall, based on forensic analysis and farmer testimony (Goldstein,
1997). The mortality incidents were highly publicized in the scientific and lay news
media, resulting in the establishment of an international working group whose
function it was to resolve potential future conflicts between agricultural production
and wildlife habitat use prior to the 1996–97 austral summer season. University
scientists, agrochemical representatives, conservation activists, and government per-
sonnel from Argentina, the U.S., and Canada joined together to designate an MCP-
free zone in the area of previous Swainson’s hawk mortality.
During 1996, use of MCP in alfalfa or as a grasshopper control agent was made
illegal in Argentina. An ecotoxicology program was initiated, with field and labo-
ratory training for government agents, students, and veterinarians living in the

pampas. Grassroots campaigns described grasshopper-eating hawks as allies to farm-
ers during the time when they were required to transition from MCP to another
chemical. The OP dimethoate and the synthetic pyrethroid cypermethrin were most
frequently chosen as chemical alternatives. With the successful removal of MCP
from this zone, hawk mortality was completely eliminated.
Agricultural Practices in Coffee Agroecosystems
Coffee (Coffea arabica) originated in Africa and was introduced to Latin Amer-
ica in the early 18th century by the Dutch. Nearly one third of the world coffee
now comes from Latin America where it is the leading agricultural commodity for
many countries and the leading source of foreign exchange. In all, 44% of the
permanent cropland is now coffee, including 750,000 ha in Central America (Per-
fecto et al., 1996).
Coffee is a shade-tolerant species and was traditionally grown under the canopy
of taller trees, often native species. Coffee in the traditional system was allowed to
grow fairly tall (3 to 5 m) under a 60 to 90% cover of shade. Plants were grown at
a relatively low density (1000 to 2000/ha), took 4 to 6 years to first harvest, and had
a life span of over 30 years. Soil erosion was low and there was little need for
agrochemical use (Perfecto et al., 1996). Several factors influenced the shift to a
more-intensified approach to cultivation, called sun coffee. First, the spread of coffee
leaf rust to Latin America caused phytopathologists to reason that the problem would
be minimized if coffee were grown in the sun as, therefore, less moisture would
accumulate on the leaves. This led to the development of more densely planted,
high-yield varieties that would produce up to four times the kilograms per hectare
of traditional plantations. Sun coffee is kept shorter (2 to 3 m) and planted at densities
of 3000 to 10,000/ha. Time to first harvest is shorter (3 to 4 years), but plantation
life span is less (12 to 15 years). In addition, there is a greater input of agricultural
chemicals and a higher likelihood of erosion. The high cost of inputs, however, made
sun coffee nearly 50% more expensive than shade coffee (Perfecto et al., 1996).
This does not include the environmental cost of sun coffee production. Nevertheless,
sun coffee has spread throughout the region and now is the most common practice
in Colombia (60% of all production; Perfecto et al., 1996).
Concern over neotropical migratory birds (NTMBs) has refocused attention on
shade coffee. Wunderle and Waide (1993) surveyed overwintering neotropical
migrants in the Bahamas and the Greater Antilles and observed that shade coffee
plantations provided habitat for species normally restricted to forests. Russell Green-
berg of the Smithsonian Migratory Bird Center and colleagues have conducted
several studies on the levels of biodiversity, including neotropical migrants, that are
supported in sun vs. shade plantations. As a generalization, shade coffee supports
more biodiversity than sun coffee; however, there is a great deal of variation among
types of shade coffee that merits examination.
Greenberg et al. (1997) did a comparison of bird species composition in frag-
ments of forest, matorral (second-growth shrub land), and three types of coffee
plantations: sun coffee, shade coffee with a Gliricidia sepium overstory, and shade

coffee with a canopy of several species of the genus Inga. Both Gliricidia and Inga
and nitrogen-fixing legumes are commonly planted in shade coffee plantations in
Latin America. Forest remnants had the highest richness, followed by the matorral
and Inga sites. Sun coffee and Gliricidia had the lowest species richness, this in
spite of the fact that the Gliricidia shade plantations were at a lower elevation with
a potentially larger pool of species to draw from. This suggests that shade coffee
alone can support somewhat higher richness than sun coffee, but the canopy species
is important. When the canopy is composed of a single species, the gain in richness
approaches early second growth, but is still less than forest fragments and far less
than undisturbed forest. Greenberg recommends that plantations should be as struc-
turally diverse as possible and a mixed agroforestry system could provide this kind
of habitat.
There are data to suggest that shade coffee provides habitat for other species as
well. Estrada and colleagues (1993; 1994) found that bat species richness in shade
coffee plantations in Veracruz, Mexico was higher than in adjacent agricultural fields
but lower than in forest; the same was true for terrestrial mammals. Biodiversity of
other groups of vertebrates and invertebrates is higher as well (Perfecto et al., 1996).
Several conservation organizations are now promoting shade coffee and mixed
agroforestry systems of production as more sustainable and environmentally benign.
The Rainforest Alliance has launched an ECO-OK certification program for coffee
to support more sustainable practices (Wille, 1994). Conservation International has
a similar Sustainable Coffee Initiative with sites in Mexico, Guatemala, Colombia,
and Peru which promotes shade coffee and organic techniques (Greenberg, 1996).
Trees as Row Crops: Plantation Forestry and Wildlife
Currently in the U.S., approximately 490 million acres of land are used for
commercial timber production (American Forest and Paper Association, AF&PA,
1996). The majority is held by private individuals (59%), with the remainder held
by the forestry industry, national forests, and other public agencies (Powell et al.,
1992). Timber is important economically; however, the impacts of timber production
on wildlife vary and are often extensive. The process of clear-cutting and refores-
tation creates an artificial cycle of disturbance, resulting in truncated succession, a
loss of species richness, and a loss in structural and functional diversity. Native stock
is often replaced by nonnative or genetically “improved” species which results in a
loss of genetic purity or genotypes of native stock. Pesticides and herbicides are
also used with unknown, secondary consequences on the native flora and fauna (Noss
and Cooperrider, 1994).
One of the most obvious and frequently cited impacts of timber production on
biodiversity is an increase in habitat fragmentation. Habitat fragmentation simply
refers to the subdividing of a continuous habitat over time (Pickett and Thompson,
1978; Foster, 1980) and processes that affect an intact forest may be exaggerated
when the forest community is fragmented (Noss, 1983). Fragmentation of forests
also increases the proportion of edge to interior habitat as the size of the forest
decreases (Ranney et al., 1981). Numerous studies on NTMBs have cited increased

competition, nest predation, and cowbird parasitism associated with fragmentation
and increased edge habitat (Brittingham and Temple, 1983; Small and Hunter, 1988;
Yahner and Scott, 1988; Wilcove and Robinson, 1990; Rudnicky and Hunter, 1993).
Another major impact of timber production on biodiversity is the loss of the
habitat itself. Of the virgin forests in the U.S., 85% had been destroyed by 1980,
with losses of approximately 95 to 98% in the lower 48 states (Postel and Ryan,
1991). Species dependent on these forest habitats, such as the endangered red-
cockaded woodpecker (Picoides borealis) suffer when these habitats are destroyed
or altered (Yahner, 1995). The red-cockaded woodpecker has almost been eradicated
from its former range, in part, due to the destruction of its primary habitat, longleaf
pine forests. Longleaf pine forests have been reduced by approximately 98% and
are the most endangered forest type (Noss, 1989). This reduction is due to logging,
fire suppression, and replacement with faster growing, more economical loblolly
pine (Dickson et al., 1995). Other species that have experienced declines, or have
been extirpated at least partially because of logging practices, include the spotted
owl (Strix occidentalis), the red wolf (Canis niger), the wood duck (Aix sponsa),
the ivory-billed woodpecker (Campephilus principalis), and the passenger pigeon
(Ectopistes migratorius) (Bellrose, 1976; Paradiso and Nowak, 1982; Robinson and
Bolen, 1984; Gill, 1990; Block et al., 1995).
Although the majority of forest management practices appear to have a detri-
mental impact on biodiversity, some practices do benefit some species. For examples,
small clear-cuts (<10 ha) appear to benefit some songbirds and mammals which
require dense, brushy, vegetation for cover and food (Scott and Yahner, 1989; Hughes
and Fahey, 1991; Yahner, 1993). Edge does benefit some game animals, such as the
white-tailed deer (Odocoileus virginianus) (Yoakum and Dasmann, 1971), and many
game managers emphasize the creation and maintenance of edge despite its potential
negative impacts on other species.
Timber management practices have recently come under fire from various orga-
nizations and this increased concern over the impacts of timber management has
led in part to the implementation of the Sustainable Forestry Initiative (SFI) by the
AF&PA. Several priorities outlined by SFI directly emphasize biodiversity (AF&PA,
1996). As advances, innovations, and technologies are developed and implemented,
these negative effects on biodiversity may be ameliorated in the future.
CONCLUSIONS — NET EFFECTS OF AGROECOSYSTEMS
Loss of Biodiversity
There is little question that, over time, agriculture has resulted in a loss of
biodiversity. The losses have been particularly serious in areas where little natural
habitat was left among agroecosystems, such as in the central plains of the U.S. The
prairie fauna once included grizzly bears, wolves, mountain lions, elk, deer, and
millions of bison, all of which are now extinct throughout most of their former
grassland ranges. The eastern deciduous forest of the U.S. was nearly completely
deforested at the turn of the century, although remnant patches of forest remained

on hilltops and in valleys. As agriculture shifted westward, many species were able
to recolonize, regenerating forests and reducing the long-term severity of the agri-
cultural impact.
If lands are properly managed in a mosaic, losses of biodiversity need not be
permanent. Studies indicate that extensive areas of tropical forest, once thought to
be virgin, were at one time under intensive cultivation. Pristine forests in the Darien
gap in Panama were apparently subjected to over 4000 years of human disturbance,
and are probably no older than 350 years (Bush and Colinvaux, 1994). Diversity is
extremely high with no indication that this history of agricultural activity depleted
the biodiversity of the region. These forests never were subjected to extensive
deforestation, however, and this likely prevented large-scale local extinctions.
Indeed, Mellink (1991) observed that the presence of isolated farms in the San Luis
Potosi plateau of Mexico actually increased regional bird species richness.
The implementation of policies recommended under the 1996 Farm Act will
help to preserve patches of natural habitat and will facilitate the protection of residual
populations of native species. If patches are kept small and isolated, however, they
may not protect viable populations over long periods of time. Areas of native
vegetation should be as large as possible and interconnected via corridors of habitat
to maximize their effect. The extensive nature of modern agriculture is much different
than the practices of the precolonial people of the Darien region in Panama, and the
long-term consequences on wildlife populations under current management practices
will be negative.
Change in Community Structure
Agroecosystems simplify the environment. Generally, they contain fewer species

than the native flora, and they contain less diversity in foliage structure than native
ecosystems. They also contain far fewer species of invertebrates and vertebrates than
natural ecosystems. Thus, the communities of organisms associated directly with
agroecosystems represent but a tiny subset of the total biodiversity of the region,
and the community structure is also simplified. These changes affect not only species
diversity, but also the functional diversity of agroecosystems. When agroecosystems
are extensive and remaining habitat is fragmented, local extinctions that result from
fragmentation will also reduce the species and functional diversity of the region as
a whole. All of these effects have been observed and documented (McNeely et al.,
1990; Day, 1996).
Changes in community structure can result in changes in ecosystem function. A
common consequence of human activities is the extirpation of large predators. The
elimination of predator populations can allow herbivore populations to increase, thus
increasing the impact of herbivores on plant community structure and function. This
can lead to large-scale changes in community structure and function (Rasmussen,
1941; Paine, 1966). Changes in species composition can also affect community
processes in unexpected ways by altering the functional diversity of communities
(Tilman et al., 1997; Hooper and Vitousek, 1997). Thus, the effects of altered species
diversity can extend far beyond the boundaries of agroecosystems and can result in
major shifts in ecosystem function as well as composition.

Recommendations for the Mitigation of Impacts
The joint production of crops and wildlife is a relatively new concept (Howitt,
1995). Paoletti et al. (1992) present a table of choices among farming systems that
either reduce or enhance biodiversity in the regional agroecosystem. Indeed, given
the preponderance of lands under some form of human management, we must begin
to assign a larger role to preserving biodiversity in agroecosystems (Pimental et al.,
1992). In the U.S., the realization that most biodiversity exists on private lands has
led to the development of new initiatives to make conservation more attractive to
private landowners. California is a habitat mosaic of federal and private lands and
an extensive region of intensive agricultural development. California also is the most
biologically diverse state with the largest number of federally listed candidates or
endangered species in the country (Scott et al., 1995). Two thirds of these listed
species are on private lands. A federally sponsored program called Habitat Conser-
vation Plans (HCP) provides a mechanism where landowners agree to an overall
plan to protect an endangered species and its habitat in exchange for a permit to
alter some portion of the habitat in the planning area (Scott et al., 1995). HCPs can
vary in their geographic scope from a single parcel or landowner to larger areas and
multiple landowners. This mechanism was instituted first in California over a con-
troversy concerning several threatened species of butterflies. The success of this
experience led to the 1982 amendment to the Endangered Species Act, allowing
HCPs. Although the majority of these HCPs are in California, they will be imple-
mented nationwide, with approximately 40 plans approved and at least another 150
in progress (Beatley, 1995).
There are numerous efforts to reduce the use and impact of agricultural chemicals
on agroecosystems (Kendall and Lacher, 1994). The U.S. Environmental Protection
Agency has supported the creation of dialogue groups to assist in the resolution of
conflicts over agricultural chemical use and the protection of wildlife (Avian Effects
Dialogue Group, 1994) and has developed a new paradigm for the assessment of
environmental risk (U.S. Environmental Protection Agency, 1992). California has
become a world leader in the development of methods to reduce pesticide use
(Anderson, 1995). The sustainable agriculture movement also emphasizes the use
of native biota for pest control (Miller and Rossman, 1995).
The impact of agricultural activities on tropical wildlife is of growing concern.
Economic factors strongly influence agricultural practices in the tropics (McNeely
and Norgaard, 1992). Frequently, pressures for colonization result from economic
hardship in urban areas, and the lack of technical expertise of colonists results in a
sequence of poor land-use practices from deforestation through inappropriate agri-
culture to abandoned pasture land. This land-use succession has been referred to as
“nutrient mining” (Southgate and Clark, 1993). Southgate and Clark (1993) make
the point that farmers and ranchers in countries where crop and livestock yields have
improved seldom encroach on natural habitats. In countries with poor yields and
increasing populations, new areas are continually being cleared. Programs that
increase yields effectively buy time for the implementation of population control.
However, most donor and foreign aid agencies are currently reducing support of

agricultural development programs. Protectionism in developed countries also inhib-
its the transfer of technologies to developing countries, inhibiting their competitive-
ness. Some environmental groups unwittingly contribute to this process by ignoring
or downplaying agricultural development and emphasizing conservation efforts only,
which can exacerbate rather than mitigate pressures on protected areas. The dynamics
of land use on a mosaic landscape are complex and interrelated; it is not possible
to view wildlife conservation and agricultural development as independent processes.
Agricultural development projects funded by international aid agencies must address
conservation and the mitigation of environmental impacts, and conservation projects
must look more closely at the interrelationship between agricultural productivity and
deforestation.
Public awareness of the impacts of agroecosystems on biodiversity in general,
and wildlife in particular, has led to the development of new laws and regulations
in countries throughout the world. There is a new appreciation of the concept of the
management of the whole landscape for multiple uses, and a better understanding
of the interrelations among landscape units. Agriculturists and conservationists alike
are coming to an agreement that the old way of doing things is no longer the best
way, considering the diverse expectations of a more-educated, globally connected
populace.
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CHAPTER 10
Natural Systems Agriculture
Jon K. Piper
CONTENTS
Introduction: Environmental Problems Associated with Modern

Agriculture
The Prairie Model
An Agriculture Modeled on the Prairie Ecosystem
Elements of the Prairie Model
Perennials as Grains
Species Diversity
Research Agenda and Findings That Support the Model
Question 1: Can a Perennial Grain Yield As Well As an Annual Grain?
Question 2: Can a Perennial Polyculture Overyield?
Question 3: Can a Perennial Polyculture Provide Its Own Nitrogen
Fertility?
Question 4: Can Perennial Polyculture Manage Weeds, Herbivorous
Insects, and Plant Pathogens?
Weeds
Insect Pests
Plant Disease
Community Assembly
Concluding Remarks
Acknowledgment
References

INTRODUCTION: ENVIRONMENTAL PROBLEMS ASSOCIATED
WITH MODERN AGRICULTURE
“One Kansas Farmer Feeds 101 People and You,” proclaims a billboard alongside
Interstate 135, near Salina, KS. Modern agriculture has been overwhelmingly suc-
cessful in terms of output per farmer, acre, or hour worked. Agricultural productivity
has steadily increased as a result of technological advances in machinery, fertilizer,
and pesticides coupled with the intensive use of plant genetic diversity to improve
yield through plant breeding. For example, yields of corn and sorghum increased
severalfold in the U.S. between the 1930s and the 1980s (Jordan et al., 1986).
In terms of return on labor, industrial agriculture based on monocultures of
annual grains is unquestionably a highly productive form of food and feed produc-
tion. This productivity has arisen largely through simplifying agroecosystems into
monocultures and tailoring them to maximize yield. In the process, however, many
of the links between organisms and the soil that serve to regulate natural communities
are ignored or disrupted. The following account surveys some of the more severe
environmental effects deriving from large-scale monocultures.
With the publication of Silent Spring 35 years ago (Carson, 1962), the public
began to become aware of unforeseen environmental consequences of modern agri-
culture and to question or not whether increasing agricultural production alone was
a worthy goal. Three of the most obvious environmental consequences of high-
production agriculture are fossil fuel dependency (and its consequent contribution
to global warming), contamination of soil and water with toxic chemical residues,
and rates of topsoil loss that exceed the natural rates of soil formation. Additional
important consequences include the net depletion of aquifer water for irrigation and
the loss of biodiversity from crops, land races, and crop wild relatives.
A general consequence of our modern agricultural system is dependency on fossil
fuel–based energy. Pimentel et al. (1995) estimate that 10% of all energy used in
U.S. agriculture is expended to offset the losses of soil nutrients and water caused
by erosion. Over the last few decades, it has taken increasingly more fossil fuel
energy to produce a unit of grain in the U.S. (Pimentel, 1984; Cleveland, 1995), with
a recent ratio of total energy expended in agriculture (including transportion and
processing) to food energy consumed in the U.S. of about 10:1 (Lovins et al., 1995).
Another consequence resulting from decades of chemical application on agri-
cultural soils is contamination of surface waters and groundwaters by toxic chemi-
cals. Particularly troublesome are unsafe levels of nitrate derived from applied
fertilizer and residues from pesticides aimed at harmful insects, weeds, and patho-
genic fungi.
Nitrate concentrations in groundwater are strongly correlated with overlying land
use (Singh and Sekhon, 1979; Hallberg, 1986). Crops often do not take up all
nitrogen applied before it leaches below the zone of biological activity in soil. This
excess nitrogen in agricultural soils can slowly leach into deep aquifers, even years
after fertilizer application ceases.
By the early 1960s, the properties of such long-lived, low-toxicity, and bioac-
cumulating substances as DDT began to emerge (Carson, 1962). In temperate

regions, DDT has a half-life of 59 years. Once bioconcentrated in such top predators
as carnivorous fish and bald eagles, DDT sharply reduces the reproductive potential
of these species. Long-term exposure has been associated with mutagenesis and
carcinogenesis. DDT was banned in 1969; other long-lived pesticides were banned
in the 1970s. They were largely replaced with several types of short-lived, acutely
toxic compounds (e.g., organophosphates). Residues of many of these subtances are
still present all over the planet. Concentration of DDE (a DDT metabolite) is
increasing in some Great Lakes and Arctic species (Hileman, 1994).
It was not until 1979 that routine agricultural use of the new generation of
pesticides was linked to groundwater contamination. Researchers discovered that
some of these apparently short-lived, unstable compounds can become extremely
persistent once below the soil biologically active zone, where the usual biological
degradation does not occur (Zaki et al., 1982; Cohen et al., 1995). Just as with
nitrate, complete cessation of pesticide use would not immediately halt the increasing
presence of pesticides in groundwater.
Many of these chemicals are threats to human health, especially among farm-
workers who are exposed directly and rural families dependent upon drinking water
from wells. Accumulating epidemiological evidence suggests that agricultural chem-
icals are associated with increased risks of many types of cancers (Blair et al., 1992;
Zahm and Blair, 1992).
An additional unintended consequence of widespread and constant pesticide
application is evolution of pesticide resistance in target organisms. The result is a
need for higher application rates of some pesticides as well as continuous research
to develop new substances to control the targeted pests. This phenomenon has been
termed the pesticide treadmill; we work harder and harder to stay in the same place
but with ever-increasing costs to environmental quality.
Despite the enormous problems presented by chemical contamination and fossil
fuel dependency, the most serious problem for the long-term sustainability of agri-
culture is soil loss. Soil erosion is the primary conservation problem on much of
U.S. cultivated cropland, and occurs mostly during short intervals of heavy rain or
high wind and when the surface is not protected by a mulch or crop canopy (Larson
et al., 1997). During the last few decades, about one third of the world arable land
area has been lost through soil erosion, and this loss continues at an estimated annual
rate exceeding 10 million ha/year (Pimentel et al., 1995). On average, soil on about
90% of U.S. cropland is being lost faster than it is being formed. Because the effects
of erosion on some soil physical attributes are irreversible, erosion rates alone may
not be good indicators of soil degradation and, consequently, soil quality can decline
faster than the erosion rate. Once virgin soil is cultivated, organic carbon rapidly
decreases (Campbell and Souster, 1982), large pores crucial for soil function are
destroyed, changes in some physical properties increase the rate of erosion, rates of
nutrient leaching can increase (Blank and Fosberg, 1989), and populations of such
beneficial invertebrates as earthworms decline (Edwards and Lofty, 1975). Prairie
soils can lose 30 to 60% of their organic carbon, 30 to 40% of nitrogen, and up to
25% of phosphorus from the A horizon after only a few decades of cultivation
(Anderson and Coleman, 1985; Schoenau et al., 1989; Woods, 1989). Many of the

consequences of soil degradation, such as reduced crop productivity, have been offset
to by improvements in fertilizer and irrigation technology and the development of
new, higher-yielding varieties.
The industrialization of agriculture, typified by widespread annual monocultures,
has led to such profound problems as soil loss, loss of genetic diversity in cultivars,
fossil fuel dependency, depletion and contamination of water supplies, pesticide
poisoning of farmworkers and nontarget wild species, and development of pesticide
resistance in pests. Modern agricultural methods, while highly productive in the
short run, are sustainable only as long as topsoil is intact, fossil fuel supplies are
affordable, and effective pesticides are available. This may be justified when fossil
fuels are cheap and environmental costs can be ignored, but such practices make us
vulnerable over the long run.
Definitions of sustainability abound. In view of the issues listed above, a sus-
tainable agriculture for the Great Plains should address simultaneously several key
environmental problems of modern agriculture. It should feature reduced or elimi-
nated soil erosion, efficient use of land area and soil nutrients, improved water use
efficiency, reduced reliance on synthetic nitrogen fertilizer, decreased risk of pest
and disease epidemics, effective chemical-free weed management, reduced fossil
energy requirements, reduced chemical contamination of soil and water, and the
opportunity for farmers to hedge their bets among several agricultural products. A
good working definition of sustainable agriculture is one that includes grain produc-
tion with (1) no chemical contamination of the environment (via pesticides or
fertilizers), (2) no dependence on nonrenewables (e.g., fossil fuel, fossil water), and
(3) no net soil loss. This working definition is limited in that it leaves out such
important considerations as sociology, economy, and justice. But it provides a begin-
ning point for a biological research agenda.
By using this definition of sustainability, what type of system could simulta-
neously satisfy all three criteria? Natural grassland ecosystems provide appropriate
models of long-term sustainability because they run on sunlight and rainfall, resist
pests, weeds, and disease epidemics, and most importantly because they do not lose
soil beyond the natural rate of formation. Some of these aspects are explored in
depth in the following sections.
THE PRAIRIE MODEL
Natural grassland ecosystems may represent our best benchmarks for sustain-
ability. Prairies (1) protect the soil from erosion, (2) provide their own nitrogen
fertility requirements through the activities of both free-living and symbiotic nitro-
gen-fixing organisms, (3) avoid devastation by weedy invaders, insect pests, and
plant diseases, and (4) run on sunlight and available precipitation (Table 1).
The vegetation structure of prairies has two important general characteristics that
contribute to sustainability: the perennial plant growth habit and diversity. Prairies
are composed primarily of herbaceous perennial plants growing in diverse arrays.
The perennial roots and canopies of prairie plants provide many benefits. These
include (1) topsoil protection from wind and water erosion, (2) improved soil quality

Table 1 Comparison between Conventional, Industrial
Agricultural Systems and Native Prairie Ecosystems for
Some Factors that Contribute to Sustainability
Factor Industrial Agriculture Native Prairie
Fragility High Low
Resilience Low High
Biodiversity Low High
Potential for nutrient loss High Low
Connectance (biotic Low High
interdependence)
Energy sources Solar, fossil fuel Solar
Nutrient sources From fertilizers Locally derived,
recycled
Adapted from J. D. Soule and J. K. Piper, Farming in Nature’s Image,
Island Press, Washington, D.C., 1992. With permission.
with time, (3) restoration of original soil structure and function following distur-
bance, (4) biodiversity of soil-dwelling organisms, (5) resistance to weed establish-
ment, and (6) stable populations of beneficial insects.
Several studies have demonstrated that the reestablishment of perennial cover
on retired cropland can reduce soil erosion while restoring soil quality. The greater
root biomass associated with perennial grasses (Richter et al., 1990) gives carbon
inputs into the soil that can be several times greater than those into cultivated soils
(Anderson and Coleman, 1985; Buyanovsky et al., 1987; McConnell and Quinn,
1988) and reduces rates of nutrient leaching relative to annual crops (Paustian et al.,
1990). Active soil organic matter, available nutrients, water-stable aggregates, and
polysaccharide content may recover under perennial grasses fairly quickly (Jastrow,
1987; McConnell and Quinn, 1988; Gebhart et al., 1994; Burke et al., 1995).
The benefits of a perennial cover were recognized by the authors of the U.S.
Conservation Reserve Program (CRP), authorized by Title XII of the 1985 Food
Security Act. This program redirected monetary resources and human efforts toward
soil conservation and indirectly toward control of agricultural non-point-source
pollution. It was designed to protect the most vulnerable U.S. cropland, with a goal
to shift 16 to 18 million ha of highly erodible land from annual crop production to
perennial vegetation for 10 years. Overall, the program keeps about 595 million t
of soil from eroding into U.S. streams and rivers annually, equivalent to a 21%
reduction in erosion on cropland (Bjerke, 1991).
The second characteristic of prairies that contributes to sustainability is plant
biodiversity. Benefits of plant biodiversity include (1) nitrogen supplied by legumes,
(2) management of herbivorous insects and some plant diseases, (3) soil biodiversity,
and (4) ecosystem stability.
Legumes play a critical role in supplying nitrogen to most natural ecosystems.
Over periods of several years, perennial legumes can increase the concentrations of
both carbon and nitrogen in the soil, as well as influence the size and activity of the
microbial community (Berg, 1990; Halvorson et al., 1991). Similarly, legumes are
important in providing nitrogen within many pastures as well as multiple cropping
systems (Davis et al., 1986; Mallarino et al., 1990). Studies have consistently shown

higher dry matter yields in grass/legume mixtures than in grass monocultures (e.g.,
Barnett and Posler, 1983; Posler et al., 1993).
Biodiversity also plays an important role in pest regulation. The presence of non-
host plant species can reduce insect density by interfering chemically or visually
with host-finding behavior and thus colonization, feeding efficiency, movement
among host individuals, and mate finding (Bach, 1980; Risch, 1981; Andow, 1990;
Bottenberg and Irwin, 1992a; Coll and Bottrell, 1994). Moreover, reduced suitability
of the microhabitat can reduce insect tenure time, oviposition, and larval survival,
and can increase emigration rate (Tukahira and Coaker, 1982; Kareiva, 1985; Elm-
strom et al., 1988). In some cases, diverse stands provide a more favorable habitat
for parasitoids and predators, leading to reduced levels of insect herbivores (Letour-
neau and Altieri, 1983; Letourneau, 1987). The weight of published evidence sug-
gests that reduced resource concentration, rather than increased numbers of natural
enemies, accounts for most of the observed herbivore reductions within polyculture
(Andow, 1991).
Similarly, numerous studies have shown benefits of plant species diversity in
minimizing certain plant diseases (Burdon, 1987), particularly those diseases vec-
tored by insects (Zitter and Simons, 1980). Establishing host plants within diversified
stands can reduce insect landing rate, and thus initial colonization of the plot, by
interfering chemically or visually with host-finding behavior (Irwin and Kampmeier,
1989; Bottenberg and Irwin, 1992a,b). This can, in turn, lead to reduced levels of
disease in mixtures relative to monoculture (Power, 1987; Allen, 1989; Bottenberg
and Irwin, 1992c). Plant species diversity can thus lower the rate of pathogen
transmission among individual host plants.
Besides these benefits, biodiversity can beget biodiversity. For example, Miller
and Jastrow (1993) noted a significant relationship between underground fungal and
floristic species richness in prairie restorations. Diversity of soil organisms can have
profound effects on plant mycorrhizal associations, nutrient-uptake ability, and nutri-
ent retention and cycling.
Biodiversity can have ecosystem-level benefits, too. Several experimental studies
have demonstrated that species-rich communities are more resilient and more effi-
cient at using resources than species-poor communities (Naeem et al., 1994).
McNaughton (1977; 1985) conducted experiments involving the grasslands of
Serengeti National Park, Tanzania. Areas of roughly 16 m2 with different diversities
were marked; then exclosures were fenced to prevent grazing by migrant herds of
zebra, gazelles, and wildebeest. Grazers reduced the biomass of diverse areas by
only about 25%, whereas the less diverse areas lost about 75% of their biomass.
Nitrogen limits the number of plant species that coexist in Minnesota grasslands;
thus Tilman and Downing (1994) manipulated the number present by varying the
amount of nitrogen applied on 207 plots each of 16 m2. They started measuring
biomass in 1987. The year 1988 featured a drought which reduced biomass differ-
ently on the different plots. After the drought, the species-diverse plots produced
half their predrought biomass whereas the species-poor plots produced only about
an eighth or less. In these studies, the more diverse communities were more resistant
to change. Such studies indicate that plant biodiversity contributes directly to the

resilience of grassland communities. Tilman et al. (1996) found that, in experimental
plots of perennial grassland plant species, community productivity increased with
plant biodiversity. Moreover, soil available nitrogen was used more completely in
the more diverse plots, leading to less leaching potential.
Finally, resistance to invasion is another collective attribute of complex commu-
nities (Case, 1990; Drake, 1991). This property is important for the ability of a
community to resist weeds and other exotic organisms.
AN AGRICULTURE MODELED ON THE PRAIRIE ECOSYSTEM
Elements of the Prairie Model
Remnant plant communities of the North American prairie are persistent biotic
assemblages in which complex webs of interdependent plants, animals, and microbes
garner, retain, and recycle critical nutrients, and protect the soil. As such they provide
our best models of the types of communities needed to restore sustainable diversity
to compromised ecosystems. Such diverse species assemblages, whose composition
varies across soil types, tend to retain species, resist invasion by exotics, and are
resilient during short-term climatic variation. Agricultural systems modeled on nat-
ural grassland ecosystems would comprise diverse plantings of perennial species
that would prevent soil loss, provide much of their own nitrogen requirement via
symbiotic nitrogen fixation, and resist invasion by weeds as well as outbreaks of
insect pests and plant pathogens. They would be structural and functional analogs
of prairie plant communities, composed predominantly of representatives from four
major plant guilds: perennial C3 and C4 grasses, nitrogen-fixing species (primarily
legumes), and composites (Asteraceae). These functional groups include the majority
of prairie vegetation (Kindscher and Wells, 1995; Piper, 1995).
A major objective of research toward a sustainable agriculture is to develop
innovative methods of production that minimize negative environmental impacts.
The CRP, although very successful in terms of soil preservation, is expensive ($1.8
billion/year) (Osborn, 1993) and provides no edible product from the idled land.
Hence, the goal of the Land Institute is to develop polycultures of perennial grains
that protect the soil and provide the restorative properties of a perennial cover while
yielding significant amounts of edible grain. Grain-producing mixtures of perennial
grasses, legumes, and composites (e.g., sunflower species) would mimic the vege-
tation structure and sustainable function of native grasslands in some fundamental
ways. Species composition of such perennial mixtures would vary geographically
and with soil type. Several promising candidates for a perennial grain agriculture
have been identified and evaluated (Wagoner, 1990; Soule and Piper, 1992). Poten-
tially, the sustainable features of such an agriculture include improved soil retention
and health, more efficient use of land area, lower fossil fuel dependence, diversity
within and between crops to reduce vulnerability to pest and pathogen outbreaks,
and greater on-farm predictability and flexibility. Because approximately 20% of
U.S. on-farm energy usage is associated with traction (Lovins et al., 1995), perennial

grain agriculture, by reducing seedbed preparation and cultivation, application of
synthetic chemicals, and irrigation will also translate into savings in energy and
materials costs for farmers.
Hence, elements of the prairie model to mimic a natural systems agriculture are
(1) herbaceous perennials as grains and (2) species grown in diverse fields. The
working model, then, comprises several perennial grain species representing four
functional groups (i.e., C4 grasses, C3 grasses, nitrogen-fixing legumes, and com-
posites) planted together.
To design persistent and diverse prairie-like grain fields, one must be cognizant
of the broad similarity of grassland communities as well as the details of their
differences. Surveys of locally and regionally occurring species and their local
distributions (e.g., Piper, 1995) suggest the types of species likely to participate.
First, there are some broad rules that hold across locations (e.g., representation by
each of the four major guilds). Second, although one cannot predict perfectly the
composition of the final successful community, the general structure of natural
communities supported on different soil types gives clues to the types of species to
emphasize in the mix (e.g., nitrogen fixers on poor soils). Third, because occasional
extreme years can limit diversity considerably (Tilman and El Haddi, 1992), high
biodiversity should enable a community to weather better the wide precipitation
swings that characterize continental climates.
Perennials as Grains
The development of perennial seed crops for agronomic mixtures consists of

two interrelated efforts. The first is the breeding of new perennial grains. This can
involve the domestication of currently wild species as well as the improvement of
wide crosses between annual grain crops and their perennial relatives. The second
area comprises long-term studies of intercrop compatibility within diversified plant-
ings. This work involves studies to discern beneficial and inhibitory crop interactions,
growth and seed yield patterns, and effects on soil quality.
Examples of the first approach, the domestication of wild perennials, are exper-
iments with the cool-season grasses mammoth wildrye and intermediate wheatgrass,
the warm-season grass eastern gamagrass, the legume Illinois bundleflower, and
Maximilian sunflower. Promising examples of the second approach, the development
of perennial grains via wide crosses between annual grains and wild congeners,
include studies with hybrid grain sorghum (Sorghum bicolor × S. halepense) and
“Permontra” hybrid perennial rye (Secale cereale × S. montanum).
An obvious consideration before any new crop is adopted is yield. Seed yields
of any new perennial grains need to be sufficiently high and stable to make their
adoption by farmers compelling. A second consideration is the possible loss of long-
term viability as a species is selected for higher seed yield. Studies of several
perennial species (Reekie and Bazzaz, 1987; Horvitz and Schemske, 1988; Piper,
1992; Piper and Kulakow, 1994), however, have indicated that, within the ranges
investigated, there are no strict “trade-offs” between increased seed yield, vegetative
growth, likelihood of future reproduction, or survivorship. Jackson and Dewald
(1994) demonstrated conclusively that a severalfold seed yield increase in eastern

gamagrass was not associated with a decline in plant growth or survivorship. Sim-
ilarly, there was no apparent trade-off between higher seed yield and rhizome
production in crosses between annual and perennial sorghum species (Piper and
Kulakow, 1994). Such results hold promise for research to increase seed yield
without losing the perennial nature of a species.
Species Diversity
Intercropping, the simultaneous raising of different crops in the same place,

makes use of species that complement one another spatially or seasonally. Relative
to monocultures, intercropped systems can display more efficient use of land, labor,
or resources, increased yield, and reduced loss to insects, diseases, and weeds
(Francis, 1986; Vandermeer, 1989). Overyielding, a yield advantage in mixture
relative to monoculture, can occur when interspecific competition in a mixture is
less intense than intraspecific competition or where plant species enhance the growth
of one another. Many factors can lead to overyielding. Crops may be released from
competition for light by having different light requirements or differences in archi-
tecture that minimize shading. Roots of different species may explore different soil
layers, or crop species may have complementary nutrient requirements or uptake
abilities. Differences in seasonal period of nutrient uptake among crops can also
promote overyielding. Intercrops may be more productive than monocultures crops
for improving soil fertility, controlling soil erosion, lowering risk of total crop failure,
and decreasing crop losses to insects and diseases. Thus, intercropping may satisfy
several crop production goals simultaneously.
One difficulty facing the plant breeder is that it may be difficult to predict from
its performance in monoculture how a crop will behave in polyculture. For example,
some plants change their root architecture or patterns of nutrient uptake when grown
in association with different species (Goodman and Collison, 1982; Jastrow and
Miller, 1993). Shorter plants that are vigorous in monoculture may be shaded out
by taller neighbors in polyculture. Thus, selection of varieties for use in perennial
polyculture is inherently more complex than selection of varieties for monocultures
due to interactions between variety and cropping system that may be unpredictable.
So, in addition to the traits needed by any viable crop (e.g., adaptation to the growing
environment, tolerance of or resistance to prevailing insect or disease organisms,
and reasonably high and stable yield potential), scientists breeding crops for poly-
culture must also select for or against competitive ability, shade tolerance, and
modifications to plant architecture that allow coexistence (Smith and Francis, 1986).
Hence, evaluation of species interactions is critical in designing a breeding program
for polycultures. Moreover, in perennial systems the outcomes of species interactions
may differ in different years (e.g., Barker and Piper, 1995).
Mechanisms that reduce overlap in resource demand among coexisting species
usually involve differences in location and timing of resource use. Roots of neigh-
boring species may explore different soil layers or a requirement might be met by
different resources. Mixtures of legumes with nonlegumes frequently demonstrate
yield advantages over monocultures because the two species are tapping different
N sources which minimizes competition for this nutrient. Similarly, intercrops may

be released from competition for light, and show greater overall productivity, if
canopies of component crops occupy different vertical layers (Davis et al., 1984;
Clark and Francis, 1985). Differences in length of the growing period or in the
seasonal periods of nutrient uptake among crops (e.g., Piper, 1993a) can also reduce
direct competition and thus promote overyielding (Francis et al., 1982; Smith and
Francis, 1986).
Alternatively, a plant may benefit its neighbors by providing cover (Vandermeer,
1980), nitrogen (Wagmare and Singh, 1984), pest or disease protection (Risch et al.,
1983; Burdon, 1987), protection from desiccating winds (Radke and Hagstrom,
1976), physical support against lodging (references in Trenbath, 1976), enhancement
of mycorrhizal associations (Jastrow and Miller, 1993), or by attracting pollinators
(Rathcke, 1984). Indirect facilitation can occur where one species “traps” nutrients
that would otherwise leach or be lost from the system, and which later become
available to other species (e.g., Agamathu and Broughton, 1985).
The Land Institute’s ongoing research agenda in Natural Systems Agriculture
revolves around four basic agronomic questions:
1. Can a perennial grain yield as well as an annual grain crop?

2. Can a perennial grain polyculture overyield?
3. Can a perennial grain polyculture provide its own nitrogen fertility?
4. Can a perennial grain polyculture manage weeds, insect pests, and plant pathogens?
RESEARCH AGENDA AND FINDINGS THAT

SUPPORT THE MODEL
Question 1: Can a Perennial Grain Yield As Well As an

Annual Grain?
Work at the Land Institute to domesticate perennial grains began in 1978 with
an inventory of nearly 300 herbaceous perennial species for their suitability to the
environment of central Kansas and their promise as seed crops. A second inventory
examined the agronomic potential in 4300 accessions of perennial grass species
within the C3 genera Bromus, Festuca, Lolium, Agropyron, and Elymus (Leymus).
From these inventories, a handful of perennial species was chosen for exploring the
principles of perennial grain agriculture.
Eastern gamagrass (Tripsacum dactyloides [L.] L.) is a large C4 bunchgrass native
from the southeastern U.S. and Great Plains southward to Bolivia and Paraguay
(Great Plains Flora Association, 1986). A relative of maize (de Wet and Harlan,
1978), eastern gamagrass has long been recognized as a nutritious and highly pro-
ductive forage. Because of its high-quality seed (27 to 30% protein and 7% fat,
Bargman et al., 1989) and large seed size, however, gamagrass shows much promise
also as a grain crop for consumption by people, livestock, or both. Ground seed has
baking properties similar to those of cornmeal. The major hurdle facing eastern
gamagrass as a grain crop is low seed yield (typically around 100 kg/ha, but as high
as 250 to 300 kg/ha in some material at the Land Institute, Piper, unpublished data).

Mammoth wildrye (Leymus racemosus (Lam.) Tsveler) is a rhizomatous C3 grass
native to Bulgaria, Romania, Turkey, and western parts of the former Soviet Union.
Grain of this and closely related species was reportedly eaten by Asian and European
people historically, especially in drought years when annual grain crops faltered
(Komarov, 1934). As is typical of cool-season grasses, wild rye displays most of its
growth in late autumn and early spring. Seed yield in Land Institute trials has been
as high 830 kg/ha (Piper, 1993b).
Illinois bundleflower (Desmanthus illinoensis (Michx.) MacM.) is a nitrogen-
fixing legume that forms a deep taproot in its first year. It is native to the Great
Plains, with a range extending north to Minnesota, east to Florida, and as far west
as New Mexico (Great Plains Flora Association, 1986). It grows best during warm
weather, flowering from late June onward. Small lenticular seeds are borne within
clusters of brown pods beginning in late July. Highest yields have approached 2000
kg/ha (Piper, 1993b; unpublished data). The nutritional quality of the seeds (38%
protein, 34% carbohydrate, Piper et al., 1988) suggests great potential as a grain
legume.
Wild or Maryland senna (Cassia marilandica) is a legume native to the south-
eastern region of the Great Plains. Flowering in Kansas takes place from late August
to early September, producing racemes of insect-pollinated yellow flowers that
become brown-black pods later in the fall. C. marilandica produces thick, deep
roots, but does not appear to form symbioses with nitrogen-fixing Rhizobium bac-
teria. The Land Institute has examined its year-to-year patterns of seed yield to
address the biological question of whether a herbaceous perennial can produce a
sustained, high yield. It provides a good model for studying the population dynamics
of a high-seed-yielding perennial (Piper, 1992).
Maximilian sunflower (Helianthus maximilianii Schrad.) is native to dry to moist
open prairies throughout the Great Plains. Its range extends eastward to Maine and
North Carolina, and westward to the Rocky Mountains from southern Canada to
Texas (Great Plains Flora Association, 1986). Maximum seed yields ranged from
1460 to 1840 kg/ha in Land Institute trial plots in 1996 (J. K. Piper, unpublished
data). In addition to its potential value as a food or oil seed crop (seed is 21% oil,
Thompson et al., 1981), Maximilian sunflower appears to inhibit weed growth
allelopathically, and may therefore be especially important during the establishment
phase of a perennial grain field.
Grain sorghum (Sorghum bicolor), a native of the African continent, is grown
extensively in the southern Great Plains as a seed crop for animal feed. It is weakly
perennial in tropical regions, but is killed by frost at higher latitudes. Johnsongrass
(Sorghum halepense), a weedy relative of cultivated sorghum, is in the U.S. a
troublesome weed that overwinters by production of rhizomes, fleshy underground
stems capable of winter survival. It may be feasible to convert a tetraploid variety
of grain sorghum from an annual to a perennial growth habit by combining in hybrids
good grain quality with the ability to produce winter-hardy rhizomes (Piper and
Kulakow, 1994). The ease of making this transfer will depend on the number of
genes controlling the production of rhizomes and whether or not overwintering
ability is genetically associated with poor agronomic characteristics.

Under favorable growing conditions, high yields of some perennial grasses can
range from 1500 to 2000 kg/ha (Ahring, 1964; Ensign et al., 1983; Mueller-Warrant
et al., 1994). Extrapolated seed yields ranging from 1720 to 2090 kg/ha have been
recorded for some perennial grasses and legumes in rain-fed, unfertilized plots at
the Land Institute (Piper, 1992; 1993b; Piper and Kulakow, 1994). These experi-
mental yields compare favorably with the benchmark yield for Kansas winter wheat
of 1800 lb/ac (1960 kg/ha).
Germplasm evaluations at the Land Institute indicate that genetic variability for
improvement of such traits as seed yield, loss of seed dormancy, shatter resistance,
resistance to viral diseases, and overall vigor is available. Moreover, including in a
gamagrass breeding program such favorable types as a gynomonoecious form pro-
lifica (a mutant sex form in which all florets are female and thus seed producing)
may increase seed yield severalfold (Dewald et al., 1987). Currently, the Land
Institute’s gamagrass breeding program is focusing on 23 high-yielding, disease-
resistant families to create a synthetic variety for further improvement.
Question 2: Can a Perennial Polyculture Overyield?
Competition between individual plants should be stronger in monocultures than

in mixtures because conspecific neighbors are most similar morphologically, phe-
nologically, and in nutritional requirements. If intraspecific competition is stronger
than interspecific competition or if facilitation is occurring, plants should yield
relatively better in mixture than in monoculture, resulting in overyielding. Con-
versely, if the effects of interspecific competition are greater than those of intraspe-
cific competition (as when species differ greatly in size, for example), then plants
should perform relatively better in monocultures. If the relative effects of different
species are neutral, then there is no yield advantage to polyculture.
Overyielding is common in traditional polyculture systems of Latin America,
Africa, and Asia (Francis, 1986; Vandermeer, 1989). Whether overyielding can occur
in perennial grain systems, and whether it can persist for more than 1 year, is less
explored. A few examples from experiments performed at the Land Institute support
the model.
In a study of 28 accessions of eastern gamagrass and Illinois bundleflower, a
relative yield total (RYT) of 1.19 was obtained (Muto, 1990). This translates into a
19% yield advantage in mixture relative to the monocultures. A three-species study
using eastern gamagrass, Illinois bundleflower, and mammoth wildrye produced an
RYT of 1.26 (Barker and Piper, 1995). In both of these cases, overyielding occurred
in more than 1 year.
Question 3: Can a Perennial Polyculture Provide Its

Own Nitrogen Fertility?
The growing environment of perennial systems changes annually. Typical initial

conditions include high levels of sunlight and available soil nutrients. With time,
concentrations of some available nutrients decline while root mass and shading
increase. With canopy closure, species interactions that began as competition for

soil nutrients may end as competition primarily for light, with plants exhibiting a
trade-off between tolerance of low soil resources and tolerance of shade (Tilman,
1985). Various factors, including root architecture, physiology (C3 vs. C4 photosyn-
thetic pathway), and ability to fix atmospheric nitrogen can result in differences
among species in net soil resource levels. Understanding the net effects of perennial
grains on soil nutrient status is an important consideration for long-term fertility
management of perennial stands to be grown with few or no inputs.
The ability of legumes to fix atmospheric nitrogen via symbioses with root
nodule–forming bacteria has important implications for agricultural sustainability.
It can release a plant from competition with neighbors for soil nitrogen and promote
the growth of neighbors if fixed nitrogen subsequently becomes available to them.
Nitrogen transfer from legumes to grasses may occur via leakage and excretion from
roots (Simpson, 1965), following decay of nodules and roots (Haynes, 1980), and
by direct mycorrhizal exchange (van Kessel et al., 1985; Eissenstat, 1990). Perennial
grasses may receive from 46 to 80% of their nitrogen directly from companion
legumes (Brophy et al., 1987). In agricultural systems, legume nitrogen can prove
both energy efficient and cost-effective (Mallarino et al., 1990; Posler et al., 1993).
Ideally, in a sustainable agriculture based on perennial grain mixtures, a signif-
icant portion of the available nitrogen should arise from one or more leguminous
companion crops. Therefore, an important question is whether or not a perennial
polyculture can provide sufficient nitrogen fertility via legume nitrogen fixation to
compensate for removal of this nutrient in harvested seed.
Work at the Land Institute has provided some indirect evidence of a benefit of
a leguminous grain to companion species. A study monitored over several years
showed that Illinois bundleflower grows and yields equally well on high and low
nitrogen soil in years with adequate precipitation (Barker and Piper, 1995). This
suggests that this legume can compensate for low soil nitrogen without a measurable
penalty of lowered growth or seed yield. A piece of evidence is that eastern gama-
grass yields better, and its yield does not decline with time, in mixtures with
bundleflower relative to gamagrass monoculture (Piper, 1998). Finally, plots con-
taining Illinois bundleflower have higher soil nitrate concentrations 3 to 5 years after
establishment than plots containing only grasses (Piper, unpublished data).
Question 4: Can a Perennial Polyculture Manage Weeds,

Herbivorous Insects, and Plant Pathogens?
Weeds
Weeds are a major source of competition with crops for light, water, and nutrients.
Because most agricultural weeds are adapted to disturbed habitats, and can often-
times take up soil water and nutrients faster than crops, they are a chronic problem
where soils are repeatedly tilled. Thus, it is not surprising that herbicides account
for 69% (by weight) of all pesticide use in the U.S. Nearly 90 million ha, more than
half of all U.S. cropland, are treated with herbicide (Pimentel et al., 1991). Perennial
grain polycultures are likely to compete well against weeds. Two important aspects

of the crop/weed relationship in perennial polyculture are the possible synergistic
effects of polycultures on weeds and the direct competitive relationship between
perennials and weeds.
In contrast to rotational sequences of monocultures, which can manage weeds
over time, intercropping combines the weed-suppressing effects of different crops
within a single season. Crop polycultures may intercept more light, water, and
nutrients than monocultures. For example, mixtures of C3 and C4 species display
active crop growth during a greater proportion of the year, potentially eliminating
the need for herbicide application (Evers, 1983). If weed mass in polyculture rows
is less than what is predicted by weed mass in the respective monocultures, this may
indicate a synergistic effect of polycultures on weeds.
An intercrop advantage may accrue in two ways (Liebman and Dyck, 1993).
Greater crop yield may be coupled with lower weed growth in polyculture. This
may occur if there is greater resource preemption by crops in the mixture (less
available to weeds) or where there is allelopathic suppression by one or more crops.
Alternatively, the intercrop may display a yield advantage while failing to suppress
weed growth below levels observed within monocultures of the component crops.
This may result if the intercrops use resources not exploitable by weeds or where
intercrop use of resources is more efficient than in monoculture.
Studies done primarily in traditional cropping systems in the tropics have shown
that internal weed management in polycultures is often greater than in monocultures
of the respective crops (Liebman, 1988). Similarly, several temperate zone inter-
cropping systems have been shown to improve weed management in important
ways. Liebman and Dyck (1993) reviewed 51 published intercropping studies where
the main crop was intersown with a “smother” crop. In 47 cases (92.2%), weed
biomass was lower in mixture than where the main crop was grown alone. When
intercrops were composed of two or more main crops, weed biomass of the intercrop
was lower than in all of the component sole crops in 12 cases (50%), intermediate
between component crops in 10 cases (41.7%), and higher than all sole crops in
only 2 cases (8.3%).
For certain combinations, increasing crop species diversity per se may suppress
weeds. In an ongoing experiment at the Land Institute using experimental mixtures
of 4, 8, 12, and 16 perennial species (see section on Community Assembly below),
percentage cover by annual weeds in the second year was inversely related to the
diversity of perennials sown, an effect which was stronger in the third year (see
Figure 1B, later).
Second, because of their permanent canopy, deep and extensive root systems,
and vigorous regrowth in spring perennial plants should in general compete well
against weeds. Several factors affect the relative ability of crop species to suppress
weeds, however. Differences in height, canopy thickness, rooting zone, and phenol-
ogy are likely to influence crop/weed interactions. Studies of prairie restoration
typically show an initial period of dominance by weedy annuals, followed by increas-
ing dominance by an array of herbaceous perennials (e.g., Holt et al., 1995).
Effective weed control has occurred in two separate experiments at the Land
Institute. In one study, a plot containing rows planted with five densities of Maxi-
milian sunflower and a control (no sunflowers), weed biomass was significantly

reduced in the sunflower plots relative to the control (discussed in Piper, 1993b).
By the second year, a sunflower density of 3.6 plants/m2 reduced weed biomass
between rows to levels only 25 to 50% of the control. In May of the third year, weed
biomass in sunflower rows was 44% of weed biomass in the control. Here, effective
weed control was maintained across years despite changes in the weed community
from predominantly annuals in the first year to perennials by the third year.
In a second experiment that examined weed growth, a triculture comprising
wildrye, eastern gamagrass, and Illinois bundleflower at equal densities, species
combinations differed in their ability to control weeds (Piper, 1993c). Weed biomass
was consistently lowest in rows with eastern gamagrass as a component, despite
seasonal and yearly changes in species composition of the weed community. These
results point to eastern gamagrass as the primary weed controller among the three
species, probably via shading, although unmeasured underground interactions were
also likely important throughout the 3-year study.
Insect Pests
For many vegetable and grain crops, pest insect density and level of feeding
damage on host species are lower within diversified stands than in monoculture
(Risch et al., 1983; Andow, 1986; 1991; Coll and Bottrell, 1994). In an extensive
review of the literature, Andow (1986) found that 131 of 203 (64.5%) monophagous
species were reported to be less abundant in polyculture cropping systems relative
to monoculture. This phenomenon, in which host plants associated with other, non-
host plant species suffer less herbivore attack than host plants in monoculture, is
known as associational resistance (Tahvanainen and Root, 1972).
Although many of the pest management benefits obtained within diversified
annual systems may transfer to perennial polycultures, typically perennial systems
are characterized by less soil disturbance and greater year-to-year continuity of the
host species. Pests therefore have the opportunity to maintain and even increase
population density in host plant patches that are relatively stable and predictable.
Unfortunately, the long-term dynamics of insect populations within perennial
monocultures vs. polycultures remains relatively unexplored.
The Desmanthus illinoensis/Anomoea flavokansiensis association represents one
system for exploring whether perennial grain polycultures can reduce insect density
relative to monoculture. The leaf beetle A. flavokansiensis Moldenke (Coleoptera:
Chrysomelidae: Clytrinae) specializes locally on Illinois bundleflower. From mid-
June to early August adults feed on young bundleflower leaves and inflorescences.
At high densities, A. flavokansiensis can reduce seed yield and is thus an important
consideration for long-term stands that are to be grown without insecticides.
The potential to manage A. flavokansiensis via intercropping its host species
with other, non-host perennial species has been examined (Piper, 1996). Replicated
plots, comprising monocultures of Illinois bundleflower, and two- and three-species
mixtures of bundleflower with the eastern gamagrass and mammoth wildrye, were
established at two sites. Over a 5-year period, the way that the plant species diversity
affects A. flavokansiensis density on individual host plants and how beetle density
changes with time in perennial stands were examined.

Insects were counted frequently from mid-June to early August. In the first 3
years, beetle density was generally low (<1 per plant), and did not differ among
treatments. In the fourth year, however, beetle density peaked at 15 and 25 insects
per plant at the two sites, and was highest within bundleflower monoculture for most
dates. In year 5, density was again low, but tended to remain higher in monoculture
at one site. The results show that beetle density on Illinois bundleflower can be
reduced in polyculture and hold promise for the management of this type of insect
herbivore within perennial grain polycultures. Differences in beetle density were
probably due more to differences in resource concentration, combined with physical
barriers provided by the grasses, than an increased presence of natural enemies in
polycultures.
It is encouraging that density of this monophagous insect on its perennial host
was reduced in some instances in polyculture even 4 and 5 years following estab-
lishment. Monitoring of the year-to-year population dynamics of this insect may
predict how similar specialist herbivores will react to plant species diversity in
perennial grain mixtures.
Plant Disease
A potential hazard confronting stands of perennial grains is the establishment

and long-term residency of systemic diseases. Perennials may be capable of storing
inoculum from year to year, enabling spread of the disease throughout the field in
later years. One possibility for the management of such diseases is to grow host
plants within polycultures.
Eastern gamagrass is infected by the pathogens sugarcane mosaic virus strain
maize dwarf mosaic virus B and maize dwarf mosaic virus. These are aphid-borne,
nonpersistently transmitted polyviruses that infect and then overwinter in gamagrass
(Seifers et al., 1993). Such diseases have the potential to increase in severity with
time. Symptoms of infection vary from a general mosaic to chlorotic and necrotic
spots dispersed throughout the leaf. At high levels of severity, these diseases reduce
growth and seed yield (Piper et al., 1996).
To examine the potential for polyculture to manage levels of aphid-borne disease
in a perennial system, plots comprising monocultures of eastern gamagrass as well
as mixtures of gamagrass with two nonhost species, mammoth wildrye and Illinois
bundleflower, were established and monitored for 5 years (Piper et al., 1996). For
the first 2 years at one of two sites, disease incidence (percent of plants infected)
and severity (intensity of symptoms) were generally lower in plots in which bundle-
flower was a component. Because there were two biculture treatments in this study,
the main factor seemed to be the presence of bundleflower within a plot, not merely
lower host plant density relative to monoculture. Illinois bundleflower appears suf-
ficiently different, in terms of architecture and biochemistry, from gamagrass to
provide an effective barrier to vector movement. The results suggest that incidence
and severity of these viral diseases can be delayed in some mixes of perennial grains,
and that the effect can persist for more than 1 year.
This study showed that manipulations of host plant density and diversity may
be useful in designing cultural methods for minimizing disease intensity and spread

in the early years. Intercropping eastern gamagrass with such nonhost species as
Illinois bundleflower appears to represent a disease management strategy comple-
mentary to selecting for viral resistance.
COMMUNITY ASSEMBLY
Theoretical and experimental studies of community organization have revealed

some intriguing phenomena with implications for the development of stable peren-
nial grain polycultures. Such studies have shown that complexity is often the con-
sequence of specific events that occur during community assembly — events that
are generally undetectable from an examination of extant community patterns alone
(Post and Pimm, 1983; Gilpin, 1987; Robinson and Dickerson, 1987; Drake, 1991;
Drake et al., 1993). Complex stable communities in nature are not created as is;
they assemble by the addition of species through invasion and by the loss of species
through local extinction. Species that form important intermediate states must be
included within the appropriate time frame. The historical sequence in which species
are added to and removed from a community determines its final structure.
Community stability changes as species are progressively added to or subtracted
from model food webs (Tregonning and Roberts, 1979; Roberts and Tregonning,
1981). A principal result of this work is that most “new” complex systems are
unstable; stability comes with age. Drake (1990a) found that with unlimited invasion
opportunities communities eventually reach persistent species compositions where
none of the species “outside” the community can enter them. In addition, there may
be so-called Humpty-Dumpty effects (Drake, 1990b; Luh and Pimm, 1993), mean-
ing it may be impossible to reassemble a community solely from the species present
in the final, persistent state. This is because species present early in assembly that
were important in determining the final structure may not be present in the final
composition.
Such results suggest that mimicking the history of a natural community is a
powerful tool for constructing stable perennial polycultures. Persistent, diverse grain
fields may be created more efficiently by starting with many more than the desired
number of species, and allowing the community to “shake down” or “collapse,” than
by experimenting with many mixes of the desired numbers of species. If this pre-
diction is correct, it will confirm an important role for history in constructing stable,
complex communities.
In 1994, the Land Institute began a study to explore whether plant biodiversity
and the sequence of species introductions can contribute to the assembly of perennial
grain polycultures. Treatments consist of four incrementally diverse mixtures of
herbaceous perennial species that represent the C4 grasses, C3 grasses, nitrogen-
fixing legumes, and composites. Several of the species are potential perennial grains.
The initial seed mixtures comprise 4, 8, 12, and 16 grassland species (Table 2).
None of the species was present at the site prior to the experiment. Each lower
diversity treatment was nested within its higher diversity counterpart (see Naeem
et al., 1994). The treatments vary the size of the species pool, while keeping guild
representation constant.

Table 2 Species Composition of Four Diversity Treatments at the Land Institute
C4 Grasses C3 Grasses Legumes Composites
I Tripsacum dactyloides Leymus racemosus Desmanthus illinoensis Helianthus maximilianii
II Tripsacum dactyloides Leymus racemosus Desmanthus illinoensis Helianthus maximilianii
Sorghum bicolor × S. halepense Elymus glaucus Dalea purpurea Helianthus mollis
III Tripsacum dactyloides Leymus racemosus Desmanthus illinoensis Helianthus maximilianii
Panicum virgatum Agropyron intermedium Lotus corniculatus Ratibida pinnata
IV Tripsacum dactyloides Leymus racemosus Desmanthus illinoensis Helianthus maximilianii
Panicum virgatum Agropyron intermedium Lotus corniculatus Ratabida pinnata
Eragrostis trichodes Agropyron smithii Amorpha canescens Liatris pycnostachya

Once seeded, half the plots within each treatment are left alone to assemble
without further intervention. The other half of the plots are resown with any species
that fail to establish, or that disappear after having established initially. The first
treatment broadly corresponds to a theoretical recipe of letting excessively species-
rich systems collapse to stable end points (Tregonning and Roberts, 1979; Roberts
and Tregonning, 1981), the second to methods of allowing both invasion and local
extinction during assembly (Post and Pimm, 1983; Drake, 1990a).
During the first 4 years, plot diversity varied considerably in the 12- and 16-
species treatments, but remained steadier in the 4- and 8-species treatments (Figure
1A). Lower diversity in the fourth year for the higher diversity treatments was due
to declining numbers of weeds in these plots (Figure 1B). Plant communities of
Figure 1 (A) Species diversity and (B) percentage cover by annual species over 4 years for
four diversity treatments seeded in 1994 (n = 16 plots). For each year, symbols with
the same letter do not differ at p < 0.05.

healthy prairie are over 99% perennial species, whereas annuals constituted most of
the cover within the community assembly plots. By the second year, 1995, however,
the percentage of cover by annuals was already decreasing as treatment diversity
increased. In the fourth year, weedy cover in the most diverse treatment was ~1/10
that of the least diverse treatment (7 vs. 65%). Even field bindweed (Convolvulus
arvensis), a pernicious perennial weed, declined as treatment diversity increased
(data not shown). Of the four major plant guilds, percentage cover by perennial C3
grasses and composites increased significantly with diversity treatment, and legumes
showed a trend in this direction (Figures 2 and 3). Percentage cover by perennial
grasses and composites is on a steady upward trajectory, with overall cover by
perennial composites, in particular, increasing more than 12-fold from 1994 to 1997.
Figure 2 Percentage cover by (A) perennial C3 grasses and (B) perennial C4 grasses over
4 years for four diversity treatments seeded in 1994 (n = 16 plots). For each year,
symbols with the same letter do not differ at p < 0.05.

Figure 3 Percentage cover by (A) perennial legumes and (B) perennial composites over 4
years for four diversity treatments seeded in 1994 (n = 16 plots). For each year,
symbols with the same letter do not differ at p < 0.05.
The results show that, after just 2 or 3 years, assembly of these plots toward diverse,
perennial communities becomes apparent.
CONCLUDING REMARKS
For several years, biodiversity has been the major buzzword among conservation
biologists as measures of diversity are commonly used as indicators of the condition
of ecological systems. Such books as Olson et al. (1995) demonstrate that the concept
of diversity is beginning to find significant applications in U.S. agriculture, too.
Diversity is widely thought to be a beneficial, even essential, property of healthy

biological, social, and economic systems, and agriculture intersects the realms of
all three.
Many of the environmental problems associated with industrialized agriculture
arise from the monoculture mind-set. Perennial grain polyculture, as a component
of a diversified farming operation, could simultaneously produce significant
amounts of grain while providing the benefits of soil and water conservation
currently achieved through the CRP. It would enhance the long-term viability of
agriculture by providing food without exhausting or contaminating the natural
resource base upon which productivity depends. Perennial grain fields could provide
an alternative to the CRP, increase year-to-year predictability, and allow greater
within-season flexibility (i.e., provide grain, hay, or pasture). In exploring this
paradigm shift from annual grain monoculture to perennial grain fields as analogs
of natural grassland ecosystems, we acknowledge that the work is inherently long
term and transdisciplinary.
The grain varieties that would be grown in perennial polyculture do not yet exist.
Thus, a long-term domestication process is required. Because we may not see great
progress for a few years, such an endeavor requires great patience on the parts of
both scientists and funding agencies. Moreover, because perennial grains will be
required to maintain themselves in the field for a period of years instead of months
as with annual crops, it is necessary to incorporate multiple-year patterns of growth
and seed yield into crop development research. For example, will yields be predict-
able year to year, or will they oscillate wildly? How will interactions among crop
species in polyculture change over years? Unlike the grower of annual crops, who
has some flexibility in changing crops after each growing season, the grower of
perennial polyculture will have to project over several possibly very different grow-
ing seasons.
The work is also necessarily interdisciplinary and cooperative, not only within
the plant sciences but among the plant sciences and other sciences, as well as among
the “hard” sciences and the social sciences. It is, therefore, going to require a great
deal of cooperation among people who previously may not have interacted much
outside their areas of expertise. A research team to explore natural systems agricul-
ture would include one or more ecologists, including those who specialize in
crop–weed interactions, soils, insects, and plant pathogens. Several plant breeders
would be needed to focus on perennial grasses, legumes, and composites. The crop
development and ecological research would be aided by scientists with expertise in
biotechnology and computer modeling. An environmental historian would put the
work in context. This list is by no means exhaustive, and several additions are likely.
The work at this point remains somewhat risky both in terms of satisfying funders
seeking short-term payoffs as well as for professional advancement (Soule and Piper,
1992). As we continue to deplete our soils and contaminate the environment, how-
ever, the long-term riskiest path for agriculture may well be the one we are on at
present. Perennial grain agriculture represents a largely unexplored approach that
will provide new avenues and insights for agricultural researchers and other scien-
tists. Innovative and creative alternatives, however, are likely to provide the greatest
insights and discoveries within science.

Humanity’s utter dependence upon a thin layer of active soil is summed up in
“Albrecht’s dilemma,” named after Missouri soil scientist William A. Albrecht. To
feed ourselves, we appropriate nature’s bounty by plowing up the prairie soil,
planting crops which produce more useful food than could be obtained from the
wild, but in the process accelerate the loss of nutrients from the soil. The movement
of productive soil from the land to the sea is hastened, to our peril, by this necessary
human activity. Opie (1993) has stated that Albrecht’s dilemma is in essence an
ethical paradox facing humanity and the environment. It is the conflict between the
inevitability of soil destruction by farming and the imperative of soil protection to
ensure a continued food supply. Most approaches toward a sustainable agriculture
can at best reduce somewhat the present environmental problems created by modern
agricultural production. Natural systems agriculture, in contrast, offers the possibility
of achieving ecological restoration as a consequence of agricultural production.
Studies of experimental grassland systems are showing that diversity, productivity,
and sustainability can go together (Tilman et al., 1996). The question facing us is
whether the tragedy of soil loss will be allowed to continue or whether the natural
systems approach will be explored as a means of reconciling the two priorities of
food production and preserving our ecological capital.
A perennial polyculture of grains is really a “domesticated prairie.” It is admit-
tedly a radical departure from current agriculture. It involves new crop species,
selected under much more complex regimes than are used for crops destined for
monoculture, and new designs for crop mixtures. Novel approaches toward a sus-
tainable agriculture highlight the critical importance of maintaining biodiversity.
Efforts to safeguard wild species, and the ecosystems that contain them, are necessary
to provide humanity with the widest possible range of options for the domestication
of perennial grains. The principles discovered along the way are likely to be appli-
cable across an array of agroecosystem types. Thus, in the work to create a natural
systems agriculture much new research ground must be broken, but in the process,
it is hoped, the broken ground of the prairie will begin to be healed.
ACKNOWLEDGMENT
I thank Brian Donahue for critiquing an earlier version of this chapter.
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CHAPTER 11
Managing Agroecosystems as
Agrolandscapes: Reconnecting
Agricultural and Urban Landscapes
Gary W. Barrett, Terry A. Barrett, and John David Peles
CONTENTS
Introduction
Cultural and Historical Perspectives of the Present Agrolandscape
The Creation of an “Oxbow” Urban Area
Linkages between Agricultural and Urban Components of the Landscape
Linkages between Biodiversity and Sustainability
Toward Sustainability of Agrolandscapes
Concluding Remarks
Acknowledgment
References
INTRODUCTION
During the past decade, several interface fields of study, including agroecosystem
ecology and landscape ecology, have emerged that integrate ecological theory and
management practices within the realm of applied ecology (Barrett, 1984; 1992).
Agroecosystem ecology is based on the premise that natural ecosystems are models
for the long-term management of agriculture and on the philosophy of working with
nature rather than against it (Jackson and Piper, 1989; Barrett, 1990). Landscape
ecology considers the development and dynamics of (1) spatial heterogeneity, (2)
spatial and temporal interactions and exchanges across the landscape, (3) influences

of spatial heterogeneity on biotic and abiotic processes, and (4) the management of
spatial heterogeneity for societal benefit (Risser et al., 1984).
The management of agricultural systems has traditionally focused on the agro-
ecosystem (i.e., crop field or landscape patch), rather than on the total agrolandscape
(i.e., watershed or region in which the crop fields are elements in the landscape
matrix) level of resolution (Barrett, 1992). Increasing the crop yield has been the
main management goal (National Research Council, 1989). Policies and practices
to maximize crop yield have involved the use of increased subsidies such as for
fossil fuels, fertilizers, and pesticides (National Research Council, 1989). These
management strategies have resulted in (1) decreased crop and biotic diversity, (2)
net energy loss, (3) profit loss for farmers, and (4) extensive nonpoint pollution of
the environment (Altieri et al., 1983; National Research Council, 1989; Barrett and
Peles, 1994).
In recent years, agricultural management strategies have begun to focus on
increasing biotic (genetic, species, landscape) diversity (Barrett, 1992), on reducing
energy inputs (Odum, 1989), and on increasing food safety (see National Research
Council, 1996, for a review) rather than only on crop yield. It has become increas-
ingly clear that we cannot sustain agricultural productivity by viewing agricultural
systems independent from other landscape elements or ecological/urban systems
(i.e., we must develop a holistic agrolandscape perspective in addition to an agro-
ecosystem perspective). We argue that a landscape approach must be established in
which landscape units, such as watersheds, are managed as functional systems based
on the concept of holistic, long-term sustainability (Lowrance et al., 1986; Barrett,
1992). This holistic approach differs from a picture of the world according to
Callicott (1989) which breaks a highly integrated functional system into separate,
discrete, and functionally unrelated sets of particulars. A piecemeal or fragmented
approach permits the radical rearrangement of parts of the landscape without concern
for upsetting the functional integrity and organic unity of the whole. By definition,
and by necessity, the agrolandscape approach must integrate aesthetic, biological,
physical, and ecological factors; must couple urban (heterotrophic) with rural
(autotrophic) systems; and must establish land-use policies based on sound ecolog-
ical theory (Barrett, 1989; Elliott and Cole, 1989). Sustainability, a common theme
of many recent paradigms, is defined here as the ability to keep a system in existence
or to prevent it from falling below a given threshold of health (Barrett, 1989).
Goodland (1995) defined sustainability, as it pertains to the environment, as “main-
tenance of natural capital.”
The sustainable landscape approach, which considers agroecosystems as com-
ponents of the total landscape (Jackson and Piper, 1989), encourages the integration
of concepts such as sustainable agriculture, biotic diversity, and levels of organization
(Barrett, 1992; Barrett et al., 1997). The focus of this approach is to manage for
sustainability of the total landscape based on an understanding of how agroecosystem
units function as an integrated whole (Figure 1).
In this chapter, we provide a perspective regarding the development and inte-
gration of modern agrolandscapes based on the ratio of primary productivity (P) to
maintenance costs (R) at the agro–urban landscape scale. This perspective is
intended to provide long-term sustainability and increased biodiversity. We discuss

Figure 1 Diagram depicting the new integrative field of study termed sustainable landscape
ecology.
the importance of providing linkages between agricultural systems and urban sys-
tems and note the importance of developing land-use policies necessary to manage
for sustainability and biodiversity based on a total landscape approach.
CULTURAL AND HISTORICAL PERSPECTIVES OF THE

PRESENT AGROLANDSCAPE
The Roman writer Cicero termed what is currently considered the cultural land-
scape “a second nature” (alteram naturam). This cultural landscape, or second
nature, comprised all the elements introduced into the physical world by humankind
to make it more habitable. Hunt (1992) interprets Cicero’s phrase, a second nature,
as implying a first, or primal nature before humans invaded, altered, or augmented
the unmediated world.
Various ideologies resulting from this second nature, especially how nature
should be managed or controlled, have contributed to the present fragmented land-
scape. The evolutionary significance of the mature (model) system, including how
natural selection has resulted in the evolution of efficient mechanisms for insect pest
control, nutrient recycling, and mutualistic behavior, is often poorly understood. A
hallmark of these mature and sustainable ecological systems is also maximum
biological diversity (Moffat, 1996; Tilman et al., 1996; Tilman, 1997). Environmental
literacy must increase if societies are to develop sustainable agriculture and sustain-
able agrolandscapes (Barrett, 1992; Orr, 1992). For example, natural processes and
concepts such as pulsing, carrying capacity, natural pest control, nutrient cycling,
positive and negative feedback (cybernetics), and net primary productivity must be
understood by ecologically literate societies in order to provide a quality environment
for future generations. There exists an urgent need to understand these processes
and concepts better, and to manage agroecosystems at the agrolandscape level
(Barrett, 1992). It is now imperative to couple the heterotrophic urban environment
with the autotrophic agricultural environment if societies are to establish or manage
sustainable landscapes on a meaningful regional or global scale.

Environmental literacy also includes the aesthetic languages of diverse cultures
and histories that determine what a people traditionally considers essential and
nonessential resources within cultures. Shifting economic, social, political, or artistic
perspectives, for example, affect the definition of what is considered a resource and
what is perceived as a nonresource. The encoded messages endemic to these cultures
influence human thought in the determination of what is of value in the life of a
human being.
The cultural landscape is an integral part of the holistic agro–urban landscape
perspective. Nassauer (1995), for example, recognized the need to investigate the
relationship between cultural landscape patterns and ecological landscape processes.
The aesthetics that are intrinsic to various cultures have influenced the present
agrolandscapes. Acknowledging these relationships, including their present and
future influences, will increase dialogue among biological, physical, and social
scientists; among resource managers, landscape engineers, and urban planners; and
among scholars investigating the role of sustainability at the landscape and global
levels (Huntley et al., 1991; Lubchenco et al., 1991). The resulting interfaces among
fields of study will lead to a deeper understanding of why and of how landscape
elements (patches, corridors, and the agromatrices or urban matrices) are related to
present regional/global patterns of belief systems, an understanding necessary to
conserve biological diversity.
THE CREATION OF AN “OXBOW” URBAN AREA
Figure 2 depicts an urban environment, including the relationship of the inner

urban landscape to the outer agricultural landscape. Although much has been written
regarding the pattern and shaping of the landscape from prehistory to present day
(see review by Jellicoe and Jellicoe, 1987, for details), there exists the need to address
and quantify the concept of landscape sustainability from an energetic (solar energy
and energy subsidy) perspective. One objective of this chapter is to increase trans-
disciplinary dialogue concerning this need. Although we recognize that markets have
become increasingly global in structure and function (Brady, 1990), it appears that
management practices, for example, integrated pest management and information
processing, will be conducted on a regional basis (Elliott and Cole, 1989).
Traditionally, towns and cities were integrated in a sustainable manner (Figure
3). The town served as the marketplace for farmers to sell their goods and products
(Mumford, 1961; Hough, 1995); goods and services radiated from the city to support
the agricultural landscape, including providing cultural, educational, and social
benefits (Le Corbusier, 1987). During the early part of this century in the agricultural
Midwest, crop diversity was high (Barrett et al., 1990), as was species and habitat
diversity (Barrett and Peles, 1994). The shift from a biologically diversified and,
perhaps, a sustainable landscape to monoculture or diculture crops (especially corn
and soybean) in the rural landscape was accompanied by the development of sub-
urban areas that reduced not only the amount of arable land, but also the diversity
of wildlife habitats and cultural linkages between the inner city and the agricultural
landscape. This created what we term an oxbow city, analogous to the creation of

Figure 2 Diagram depicting urban, suburban, and exurban/agricultural systems. Solar-pow-
ered (autotrophic) patches are shown within urban and suburban (heterotrophic)
systems.
an oxbow lake when it becomes separated (physically and functionally) from a

flowing meandering stream once the stream changes its course. This isolated city
develops different functional processes (i.e., provides different services), resulting
in changes in niche and biodiversity (i.e., the inner city creates different occupations
and provides habitats for different species of flora and fauna). The integrity of the
city frequently becomes less closely related to the total watershed from which it
evolved. This developmental process is depicted in Figure 4.
LINKAGES BETWEEN AGRICULTURAL AND

URBAN COMPONENTS OF THE LANDSCAPE
Odum (1997) classified ecosystems based on the proportions of solar and fossil
fuel energy used to drive the system. Most natural ecological systems are driven
entirely by solar energy. Subsidized systems depend, to varying degrees, on the input
of subsidies such as fossil fuel energy, fertilizers, and/or pesticides. Agroecosystems,
for example, are driven by both solar energy and subsidies; urban systems depend
mainly on enormous inputs of fossil fuel subsidies (Odum, 1989).
These ecosystems may also be classified based on the ratio of energy produced
by primary productivity (P) to energy used for respiration or system maintenance
(R). Natural and agricultural ecosystems, especially during ecosystem growth and
development, represent autotrophic systems where P/R > 1. In contrast, urban areas
have increasingly become heterotrophic (P/R < 1). We define sustainable systems
as those systems or landscapes where long-term P/R ratios equal 1. During the
growth and development of autotrophic systems (i.e., during ecological succession),

Figure 3 The development of an agro–urban sustainable (P/R = 1) landscape. The town
marketplace historically was closely linked to the agricultural landscape. Sustain-
ability in the modern agro–urban landscape increasingly must be based on the
management of suburban areas (ecotones) as natural linkages between urban and
agricultural systems.
P/R decreases as biological (organic) materials accumulate (Figure 5). This results
in a balance between productivity and respiration (P/R = 1) in the climax stage of
succession (Odum, 1969). An increase in physical (inorganic) materials in urban
systems coincides with a decreasing P/R ratio (i.e., a significant increase in main-
tenance costs). Thus, the result of urban succession is a city where energy demands
greatly exceed productivity. During the past century, the large numbers of people
living in urban and suburban areas have led to increased need for food produced in
rural areas (Steinhart and Steinhart, 1974; Odum, 1989). This demographic and
cultural transition has led to increased reduction of P/R ratios in urban areas, as well
as increased subsidization of agriculture (including economic subsidies) to maximize
crop yield (National Research Council, 1989; 1996).
More recently, suburban expansion has led to increased pressure on rural land
used for agriculture (Lockeretz, 1988). A result of urban sprawl has been an increase
in the proportion of the agrolandscape occupied by heterotrophic systems. This has
serious implications regarding the conservation of biodiversity from a sustainable
agrolandscape perspective (Rookwood, 1995). In addition, urban expansion into
agricultural land has important consequences for aesthetic, social, and economic
values (Lockeretz, 1988), including the need to understand more fully how human

Figure 4 Diagram depicting the development of urban areas into oxbow cities (1 to 3) and
then, we hope, into a modern, sustainable agro–urban landscape. We argue that
the modern landscape must be increasingly based on the ecological/economic
management of suburban areas as natural linkages between urban and agricultural
systems.
values will likely define future landscape boundaries and resources, especially those
values that relate to ecosystem/landscape sustainability.
The present challenge for agrolandscape management is to minimize the infringe-
ment of urbanization on agricultural land, to restore biological diversity (genetic
niche, species, and landscape) at greater temporal and spatial scales, to establish
linkages (ecological and economic) between urban and rural (heterotrophic and
autotrophic) patch elements, and to achieve sustainable productivity (P/R = 1) at
agro–urban (regional) scales. Goals for achieving sustainable agrolandscape man-
agement should focus on (1) achieving stability regarding P/R ratios among het-
erotrophic and autotrophic systems at these scales; (2) creating both natural corridor
and human transport linkages between rural and urban systems; (3) protecting the
integrity of ecosystem/watershed processes, such as nutrient recycling and primary
productivity; and (4) establishing management policies for optimal land use within
transition suburban areas that ecologically and economically form an interface
between urban and agricultural landscape systems. As previously noted, sustainable
agriculture is based on the coupling of agricultural ecosystems with natural ecosys-
tems (Barrett et al., 1990). Here, we stress the need to integrate natural, agricultural,

Figure 5 Changes in P/R ratios during ecological (autotrophic) and urban (heterotrophic)
succession. Although P/R decreases as the amount of biological material increases
during ecological succession, the result is a mature (climax), sustainable community.
In contrast, the accumulation of physical material during urban succession frequently
results in a fragile, nonsustainable community.
and urban components if societies are to design and implement the concept of
sustainability at the agrolandscape scale (see Figure 1). This approach should simul-
taneously conserve and enhance biotic diversity at greater temporal and spatial
scales.
LINKAGES BETWEEN BIODIVERSITY AND SUSTAINABILITY
Management of agrolandscapes for sustainability both influence and is influ-

enced by biodiversity (Paoletti, 1995). Landscape planning is a process through

which the conservation and management of biodiversity can be pursued (Rook-
wood, 1995).
Turner et al. (1995) stressed that there exists a three-way interaction of biodi-
versity, ecosystem processes, and landscape dynamics at greater scales. Sustainable
agricultural practices leading to increased crop and genetic diversity have resulted
in increased agroecosystem stability (Cleveland, 1993). For example, increasing crop
diversity benefits agriculture by reducing insect pests (Altieri et al., 1983). Other
sustainable agricultural practices, such as conservation tillage, are known to increase
habitat diversity, wildlife diversity, and numbers of beneficial insect species (Barrett,
1992; McLaughlin and Mineau, 1995).
Although the importance of a landscape approach for management of biodiversity
is well recognized (Franklin, 1993), little is known regarding how biodiversity affects
landscape pattern and dynamics (Turner et al., 1995). Turner et al. (1995) have
suggested that there exists a three-way interaction among biodiversity, ecosystem
processes, and landscape dynamics. In addition, it is well documented that there
exists a reciprocal relationship between sustainability and biodiversity (Paoletti,
1995). Thus, as biodiversity and sustainability are increased by management prac-
tices at the landscape level (e.g., agrolandscape management), the resulting increase
in biodiversity will likely have important benefits concerning the conservation and
efficiency of these processes and dynamics (Culotta, 1996; Tilman et al., 1996).
It is important to recognize that, by definition, the agrolandscape approach
requires the consideration of biological diversity in the management of agroecosys-
tems (Paoletti et al., 1992). Paoletti et al. (1992) and Paoletti (1995) note that
sustainable strategies in food production in agriculture improve the existing biodi-
versity. These strategies include proper management of natural vegetation, better use
and recycling of organic residues, introduction of integrated farming systems,
reduced tillage, intercropping, crop rotation, biological pest control, and increased
number of biota involved in human food webs. McLaughlin and Mineau (1995)
point out, however, that agricultural activities such as tillage, drainage, rotation,
grazing, and extensive usage of pesticides and fertilizers have significant implications
for wild species of flora and fauna. Therefore, reduced or (no-till) farming, in contrast
to conventional tillage, benefits biological diversity in terms of maintaining wild or
native species populations.
Increased biodiversity at the landscape level (in the form of increased habitat or
agroecosystem diversity) will play a key role in protecting diversity and in providing
a linkage between urban and rural areas in our sustainable landscape approach. For
example, that an optimum balance between solar-powered and subsidized systems
in suburban areas is critical to this linkage. The success of obtaining this optimum
balance will almost certainly be enhanced by increasing the diversity of habitat types
across the total landscape. Greenways or natural corridors will also enhance the
linkage and conserve biological diversity between urban and rural areas (Little,
1990). Management of agroecosystems and agrolandscapes for sustainability will
lead to increased habitat and genetic diversity, which, in turn, will lead to increased
agroecosystem stability (Altieri et al., 1983; Cleveland, 1993). Likewise, increased
biodiversity within individual systems should also increase the diversity and stability
of these systems within the total landscape or watershed.

Agrolandscapes should also be managed to increase species diversity within
landscape patches and to increase and/or to conserve genetic material among land-
scape patches (Barrett and Bohlen, 1991). In recent years, there has been increased
emphasis on the connectivity and integration of the agricultural landscape with the
urban landscape (see Lockeretz, 1988, including a special issue of Landscape and
Urban Planning, Volume 16, for details). However, most studies at the watershed
or agrolandscape levels have failed to encompass or integrate the urban environment
into the agrolandscape concept. Since the approach to sustainable agriculture is
based on natural ecosystems serving as the model system for efficient agricultural
management (Jackson and Piper, 1989; Barrett, 1990), urban systems should also
be designed and based on natural ecological systems serving as model systems to
ensure maximum ecological and economic efficiency. Thus, sustainability and biodi-
versity share an important interrelationship that is more fully understood when
questions are addressed at the agro–urban landscape scale, and when research
designs and management strategies are based on ecological theory.
Importantly, there is growing recognition that cities need to be managed based
on the concept of sustainability (Stren et al., 1992). This approach is based on an
ecological understanding of how natural ecological systems are organized, and most
important, how they function. As with sustainable agriculture, an urban perspective
based on sustainability means working with, rather than against, nature. Recently,
there has been increased effort in urban areas to maximize the efficiency of energy
use, to increase the rate of recycling of goods and materials, and to reduce pollutants
entering the system. In addition, “green city” movements have placed emphasis on
preservation of natural areas, and on the establishment of vegetable gardens (i.e.,
solar-powered patches, Figure 2) in urban areas (Stren et al., 1992). Continued efforts
to increase solar-driven productivity, while simultaneously decreasing maintenance
costs, in urban areas will greatly enhance the sustainability of the total agro–urban
landscape. Equally important is the need to plan for (or to zone) future suburban
areas, encompassing a productivity/maintenance ratio equal to 1, if we are to achieve
regional landscape sustainability (Figure 4). This strategy must also make every
effort to increase biodiversity at the genetic, species, and landscape levels.
Lockeretz (1988) stressed the importance of protecting and creating natural
linkages between urban and rural areas. Urban greenways (Little, 1990) provide
noteworthy examples of these natural linkages. Although greenways take many
different forms, they are primarily natural areas set aside for their ecological, rec-
reational, or aesthetic value within urban areas. Greenways also provide natural
corridors for movement of wildlife species and transfer of genetic materials between
urban and rural areas.
Suburban areas also represent a vital linkage (transition or ecotone) between
urban and rural systems. Therefore, it is important to manage these areas as a
“transitional zone” between urban population centers and rural farmland (Figure 4)
with an optimum balance existing between land area devoted to natural (solar-
powered) systems and those managed as subsidized systems. Management plans
aimed at establishing a new approach to suburban development should strive to attain
an integration of autotrophic and heterotrophic systems. The success of these man-

agement plans will greatly depend on an increased ecological literacy of people living
within and in consort with future land-use policies (Orr, 1992; Barrett et al., 1997).
In recent years, numerous forms of land-use policies have been instituted to
protect agricultural land at urban-fringe areas (Luzar, 1988) and to promote sustain-
able agriculture research and education (Hess, 1991). Although these programs have
been moderately successful (Harsch, 1991; National Research Council, 1996), there
still is much to learn regarding the most effective combination of strategies for a
particular region. There is urgent need for a new integrative “greenway” strategy.
For example, the words ecology and economics are both derived from the Greek
root oikos, meaning “household” (logos meaning “study of” and nomics meaning
“management of”). Thus, ecology is the study of the household or the study of
natural systems, whereas economics is the management of the household or the
management of natural/socioeconomic systems. We must now integrate greenways,
green meaning primary production or green meaning “greenbacks” or dollars, with
ways, meaning progress in a specific direction, as a new integrative ecologic/eco-
nomic strategy.
Determining optimum land use is a difficult task (Werner, 1993). For example,
the most effective land-use strategy for urban–rural fringe areas depends on agri-
cultural, cultural, demographic, and socioeconomic characteristics of an area (Lock-
eretz, 1988). The development of appropriate management strategies at the agro-
landscape level will require the cooperation of diverse fields of study (Barrett, 1992).
This cooperation will enhance biotic diversity at the landscape level, will provide
numerous societal benefits, and will protect both natural capital (goods and services
provided by nature) and economic capital (goods and services provided by socio-
economic systems) as a long-term strategy (Costanza et al., 1997). The longer the
wait, the greater are the long-term costs regarding the loss of biodiversity, the
continued loss of soil and nutrient resources, the degradation of water quality, and
an increased cost to restore habitat quality.
TOWARD SUSTAINABILITY OF AGROLANDSCAPES
There exists the need to integrate and interface natural, agricultural, urban, and
suburban systems if humans are to manage agrolandscapes in a truly sustainable
manner. How might we approach this problem? Agricultural practices should increas-
ingly be devoted to the preservation of natural areas (e.g., forests) and to the planting
of a diversity of food and grain crops near the outer perimeter of our large cities
(i.e., should view agricultural planning in conjunction with, rather than against, the
planning of perimeter highways and other modes of transportation). Traditionally,
urban and rural landscapes were interrelated in a mutual, sustainable infrastructure.
Urban areas were frequently directly linked to the watershed or drainage basin
because of transportation needs (rivers or lakes) or geological nodes (mineral depos-
its) of significance. Urban “sprawl” into the agricultural landscape tended to decou-
ple this connectivity and sustainability. Rather than serving as an ecological/eco-
nomic barrier, the suburban patch should provide a critical link to restore habitat

patch connectivity (Fahrig and Merriam, 1985) and to promote societal sustainability
(Barrett, 1989).
Urban areas contain human and natural resources that once again must become
integrated with the agrolandscape to the extent that the city can be sustained by both
economic capital (gross national product, employment, property values) and natural
capital (primary productivity, biotic diversity, solar energy, and sludge usage as
natural fertilizers). We refer to this perspective as “dual capitalism.” Economic and
ecological capital must become integrated; otherwise the resulting oxbow urban
areas will increasingly become drained of aesthetic and natural resources. Ecosystem
processes will increasingly become disconnected and/or diminished from the vital
flow of energy, information, or biotic diversity that once connected the landscape
system as an integrated whole.
The integrity of the architecture, the efficiency of the natural processes, and the
quality of materials typically found within city boundaries (Kaplan and Kaplan,
1982) also have been reduced or degraded during this fragmentation process. Ecol-
ogists frequently note “that the ecosystem is greater than the sum of its parts.”
Perhaps the landscape level of organization is now no longer greater than the sum
of its parts. Isolated urban patches exhibit greater entropy (poverty and crime) and
require greater subsidies (financial and human) to maintain their existence (Jake and
Wilson, 1992). Isolation and fragmentation impede the attainment of ecological
literacy (Orr, 1992) necessary to understand how natural regulatory mechanisms
enhance the reestablishment of diversity (ethnic, biological, and ecological) which,
in turn, is necessary to maintain agrolandscape functional processes. It has become
increasingly critical to understand these relationships and regulatory mechanisms
more fully. To do so, greater emphasis should be placed on suburban patches in an
effort to integrate and reconnect urban and agricultural processes. Suburban “eco-
tones” provide the opportunity to help transform the total landscape unit. Suburban
areas are not marginal or isolated places, but increasingly central locations in the
contemporary world (Baumgartner, 1988). Table 1 contrasts urban, agricultural,
natural, and suburban systems and suggests that suburban areas be designed and
managed based on our understanding of how natural ecosystems are structured and
how they function.
Figure 4 depicts how this suburban area might be designed to interface with a
transportation network, including the establishment of an enriched food crop, for-
estry, and recreational diversity that function as an economic/ecological transition
area between the urban and rural landscapes. Landscape corridors, both natural
(streams and trails) and human built (highways and mass transportation), will further
enhance these linkages (Little, 1990).
Landscape corridors (linkages) manifest various configurations depending on
natural phenomena, cultural preferences, or historical development (Hough, 1990).
Corridors may connect a sequence of congruent patches. For example, corridors of
an ecological mosaic have been defined as disturbance corridors (e.g., power lines),
planted corridors (e.g., shelterbelts), regenerated corridors (e.g., fence-row succes-
sional vegetation), remnant corridors (e.g., strips of native or climax vegetation) and
resource corridors (e.g., riparian areas) (see Barrett and Bohlen, 1991, for details
regarding corridor types). These corridors also may reflect or parallel land-use

Table 1 Contrasts among a Traditional Heterotrophic Urban System, an Autotrophic
Conventional Agricultural System, a Natural Mature Ecological System, and a
Well-Planned Integrated Suburban System
Attribute Urban Agricultural Natural Suburban
Primary P/R < 1 P/R > 1 P/R = 1 P/R > 1
productivity
Nutrient source Commercial Commercial Nutrients Wastewater
sludge recycled effluents,
sludge, and
manures
Goal-driven r-selecteda r-selecteda K-selecteda K-selecteda
(yield)
Pest control Pesticides Pesticides and Natural Integrated Pest
Integrated Pest Management
Management (IPM)
(IPM)
Input energy Fossil fuels Solar and fossil Solar Solar
fuels
Biodiversity Low Low High High
Sustainability Short-term Short-term Long-term Long-term
a r-selected: favors lifestyle in which resources are diverted to high growth rate and fecundity
rather than persistence. K-selected: favors lifestyle in which high growth rate and fecundity
are reduced to direct resources to persistence under carrying capacity conditions.
policies, a mapping of sacred spaces, or the inventory/acquisition of natural

resources. Corridors often reflect, or parallel, political, social, economic, and artistic
preferences. Thus, the management of sustainable agrolandscapes and their restored
connectivity will require a better integration of aesthetic and ecological principles
and theory. This integration process is one of society’s greatest challenges for the
21st century.
CONCLUDING REMARKS
A telescopic/microscopic view of the landscape provides a mosaic of patches

(isolated and connected) that form a complex network of elements constituting our
20th-century landscape. The fragmentation of science and the proliferation of man-
agement policies indicative of the 20th century must be replaced by integrated
approaches during the 21st century if societies are to achieve the goal of sustainable
societies and to protect biotic diversity at regional and global scales. Lubchenco et
al. (1991) outlined a research initiative necessary to establish a sustainable biosphere
initiative (SBI) at greater scales. Programs or initiatives, such as the SBI, should
focus on the landscape level (including human settlements) with special attention
directed to protecting and restoring the integrity of ecosystem and landscape pro-
cesses and to conserving biotic diversity. Only with long-term land-use planning
can humans hope to protect and restore sustainable landscapes for future generations.
An agro–urban landscape perspective is important for ecologically self-sufficient
and economically cost-effective management of agricultural systems. Management
decisions based on this perspective should be aimed at achieving sustainability of

the total landscape. We define sustainability in energetic terms in which primary
productivity of the total landscape is in balance with community maintenance (i.e.,
P/R = 1). To achieve such a landscape sustainability, the rural autotrophic landscape
(P > R) must balance the heterotrophic urban–suburban landscape (P < R). Biotic
and cultural diversity must be maintained within these systems to protect and/or
conserve regulatory feedback mechanisms (Barrett et al., 1997).
Because it is a difficult task to determine optimal land use, the development of
appropriate management strategies at the agro–urban level will require the coopera-
tion of diverse fields of study. The political, social, artistic, and economic components
must become integrated with ecological theory in order to optimize land-use planning
and, consequently, determine the landscape mosaic for the 21st century (Barrett and
Peles, 1994). The integration of the ecological, cultural, and historical spheres of
knowledge should generate new theories and methodologies necessary for the appro-
priate management of these agrolandscapes. The coupling of theory with practice
must become an integral part of this integrative planning and management strategy.
The human process of orientation by another can have a significant influence on
what human beings perceive as a resource and a nonresource and, consequently, on
how humans view the total landscape. What people consider a resource changes
with purpose or intent while negotiating their environment (Hunt, 1992). This per-
ception, in turn, changes the status of other resources. For example, humans increas-
ingly must combine ecological “capital” with economic “capital” (dual capitalism)
in order to manage these systems and landscapes best for future generations.
The integration of the agricultural, urban, and natural landscapes, landscapes
that transcend cultural (i.e., intragenerational) and historical (i.e., intergenerational)
boundaries, must be more fully understood in order to optimize biological and
landscape diversity. Linking an increasing number of diverse cultural and historical
perspectives with the future design of the total landscape, through dialogue, changes
the pattern of our understanding of landscapes. As Nassauer (1995) notes, changing
perceptions change the landscape. The role of biodiversity in total landscapes, viewed
and researched based on this perspective, can best be managed within this continuing
dialogue.
Future generations will require this type of mutualistic behavior and transdisci-
plinary cooperation to manage increasingly complex regional and global landscapes.
Maintaining biodiversity is paramount to the sustainability (health) of our planet
and is by necessity transgenerational.
ACKNOWLEDGMENT
We thank Eugene P. Odum and an anonymous reviewer for providing comments

and suggestions regarding this chapter.

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CHAPTER 12
Local Management of Biodiversity in

Traditional Agroecosystems
Robert E. Rhoades and Virginia D. Nazarea
CONTENTS
Introduction
Historical Perspective on Agrobiodiversity
Strategies for In Situ Agrobiodiversity Conservation by Indigenous
Communities
Multidimensional Criteria for Selection and Maintenance of
Landraces
Comparison of Scientific/Formal Approaches to Biodiversity
Maintenance
Social, Economic, and Political Dimensions of In Situ Conservation
The Social Context of Community-Based Biodiversity Management
Global Change and Plant Genetic Resources
Pathways Toward the Future
References
INTRODUCTION
The development goal of increased food and fiber production to match the needs
of growing populations and their rising expectations may often conflict with the in
situ conservation goal of preserving plant genetic diversity (Williams, 1986; Alcorn,
1991). As broadly adapted high-yielding varieties — coupled with input packages

of irrigation, fertilizers, and pesticides — find their way into traditionally diverse,
marginal agroecosystems, the number and diversity of local landraces as well as
associated local knowledge may erode. This process is global, although its dynamic
and intensity may vary from place to place and across time (Ford-Lloyd and Jackson,
1986; Oldfield and Alcorn, 1991; Brush, 1992; Dove, 1996).
One of the premises of sustainable agriculture is that this trade-off between
increasing productivity and loss of biodiversity is not inevitable (National Research
Council, 1992; Thrupp, 1997). Precisely how the two goals are simultaneously
achieved, however, is not an insignificant research problem or an easily answered
policy issue (see Williams, 1986). Present demographic and economic global trends
require more food per unit area and unit time, but the necessary yield increases will
not be forthcoming unless sufficient biodiversity is continuously available to plant
and animal breeders. Our primary thesis is that one useful but neglected strategy to
achieve sustainable food production lies in supporting traditional in situ biodiversity
management. We argue that many local populations have historically managed
biodiversity, that the associated knowledge is valuable and irreplaceable, and that
both management practices and knowledge should be enhanced through policy and
technology initiatives.
Specifically, we address our thesis by exploring three major themes related to
indigenous management of germplasm and the potentialities for the localized cre-
ation, maintenance, and enhancement of biodiversity. First, we place biodiversity
management by traditional agroecosystems in global historical context, especially
as it relates to the major food crops. Second, based on our own research experiences
we outline some principles of in situ biodiversity maintenance within traditional,
marginalized agroecosystems and contrast these to the scientific, ex situ approaches
of formal, external input-dependent and market-linked approaches. Third, we exam-
ine the social, economic, and political dimensions of marginal communities man-
aging in situ agrobiodiversity. Finally, we conclude with some observations on future
research and action needs.
HISTORICAL PERSPECTIVE ON AGROBIODIVERSITY
Landrace-based genetic materials available for plant breeding or biotechnology

programs have already been purposely manipulated by traditional cultivators over
centuries and even millennia (Ucko and Dimbleby, 1969; Struever, 1971; Altieri
and Merrick, 1987). Although archaeological debates continue over precisely where
and how nomadic hunters and gatherers finally began the conscious planting of
seeds, roots, or tubers and thereby ushered in the “Agricultural Revolution,” the
outcome of early farmers’ efforts cannot be denied (Hobhouse, 1985; McCorriston
and Hole, 1991). The historic tendency for preindustrial agricultural communities
has been to foster and increase landrace diversity, rather than decrease it (Harlan,
1995). As long as 8 to 12 thousand years ago, “primitive” farmers had already
successfully experimented with invading wild “weedy” species in their settlement
clearances and domesticated the first crops (Harlan, 1975). Not only did prehistoric
cultivators give humanity the major food crops and animals which nourish us today,

they simultaneously created their own specialized knowledge systems about the
food, fiber, and medicinal values of thousands of plant and animal species (Schery,
1972; Fowler and Mooney, 1990). While modern science has been appreciative of
and concerned about the supply of the genetic raw material provided by farmer
curators, much less interest has been shown in the local knowledge or management
strategies which underpinned in situ landrace development in the first place (Naz-
area-Sandoval, 1990).
De Candolle (1885) and later Vavilov (1926; 1949) were the first to observe that
the density of interspecific and intraspecific variation of crop species was found in
“centers of domestication” which tended to be in the ecologically complex moun-
tainous regions or areas of marked dry–wet seasons in Africa, Asia, and Latin
America (Rhoades and Thompson, 1975). Due to a variety of causes, major ancient
civilizations — such as the Andean, Mesopotamian, Mesoamerican, Indus, and
Chinese — evolved near these centers in close association with diverse plants and
animals. In complex ecological settings under conditions of human population
expansion, the coevolution of human culture and plant populations led to a level of
people–plant interdependency so high that some modern crops — such as maize —
cannot even reproduce themselves without purposeful human intervention (Iltis,
1987). The historical and ethnographic records are rich with data on how cultural
knowledge intertwines with the biological to the degree they cannot be separated
and still maintain dynamic evolutionary-ecological systems (Nazarea, 1998a). This
detailed knowledge not only focused on production but also storage, processing,
cooking, and utilization qualities needed for the survival and rejuvenation of crops
and humans (Nazarea-Sandoval, 1992). As a result, domesticated crops can be
understood as culturally created and conceived human artifacts — valued for multiple
qualities such as utility, taste, color, shape, and symbolism (Zimmerer, 1991). Indeed,
foods or other materials derived from crops or animals are not just calories for the
human body but are integral parts of daily social and cultural lives (Brush, 1992).
The original diversification of crops in the centers of domestication was further
enhanced after the Age of Discovery when plants were transferred by explorers
between the Old and New Worlds (Hobhouse, 1985). In their new homes, migrating
plants were further manipulated by curious cultivators and horticulturists who tested
the exotic materials, selected those that did well, and then integrated them into their
local farming and gardening systems. However, problems in the transplanted plants
soon became evident. Since only a small amount of the genetic variability found in
the agroecosystems of domestication made it to the new environments, resistance
to disease and pests was often lacking and collapse under the onslaught of disease
or pests was devastatingly frequent (Rhoades, 1991). The most famous documented
case is that of the widespread potato crop failure in mid-19th-century Europe due
to late blight (Phytophtera infestans), a fungus likely introduced from the Americas.
Most of Europe had come to depend on a few varieties; in Ireland there was total
dependence on a single variety. Combined with political exploitation by the British
government, the unfortunate timing of the crop failure led to the deaths of millions
of Irish (Woodham-Smith, 1962). A few years later, the grape crop on mainland
Europe succumbed to a minute, aphidlike insect (Phylloxera vitifoliae) accidentally
introduced from wild American grape stock (Vitis labrusca) (Olmo, 1977). These

two incidents spurred a tremendous interest on the part of European scientists to
understand not only the nature of disease (plant pathology was born of these efforts),
but also the relationship among the centers of genetic origin, natural range of
variation, and disease resistance. Although neither Mendelian genetics nor the theory
of disease was understood by the 1870s, European farmers appreciated that
“renewed” seed stock from the regions where the crops originated brought bloom
back to their crops. In the case of postfamine potatoes, a single small Andean tuber
direct from South America fetched its weight in gold, thereby creating a potato seed
craze as intense as the tulip craze in Holland in earlier times (McKay, 1961).
Likewise, European and American plant scientists came to appreciate the link
between the well-being of their farmers’ crops and the genetic diversity in the
homelands of the crops themselves.
Although the importance of genetic material from the centers of diversity and
domestication remains highly appreciated by geneticists and crop scientists, there is
less awareness of the curator role of extant farmers or pastoral communities and
their knowledge which makes it possible for this valuable diversity to be maintained
in situ and passed on to the global human family. Historically, most governments
have seen marginal tribal and peasant communities as practicing a backward, prim-
itive agriculture ripe for “modernization” through information and technology (Rog-
ers, 1969). As industrial developments in Europe, North America, Japan, and the
cities of the Third World attracted wage labor from the countryside, planners and
agricultural scientists sought ways to provide cheap and abundant food for the
growing urban areas. This cheap food policy, which has intensified in the post–World
War II era, meant that the potentially productive agroecological zones (flat, fertile,
and hydrologically favorable) were to become targets of planned agricultural change
to make them more productive through genetic uniformity and mechanization of the
agroecosystem for the purpose of achieving higher yields. One outgrowth of this
simplification of the agricultural landscape was the renowned “Green Revolution,”
which combined scientific plant breeding with input packages for favorable envi-
ronments (Plucknett et al., 1987). The dramatic increases in world food supply
witnessed in the 1960s and 1970s are directly traceable to these crop improvement
programs which focused on increasing the productivity of plants though breeding
for high response to inputs such as fertilizer (Mellor and Paulino, 1986).
The role ascribed by scientists to local cultivators and their communities during
the Green Revolution was that of recipients of “technological” packages of improved
seeds, fertilizers, and other inputs, as well as infrastructure development. “Trans-
forming traditional agriculture,” as Nobel Peace Prize–winner Theodore Schultz
(1964) called the effort, was promoted as the motor for global growth and the most
efficient exit from agricultural stagnation and famine. Farmers were seen as indi-
vidual rational decision makers who only needed to be provided the necessary inputs
and knowledge by governments and scientists to get the job accomplished. Hence,
breeders made selections and crosses from advanced breeding lines derived from
landraces. These lines were tested on experiment stations or controlled farm condi-
tions, and, after a dozen or more years, these materials were released to farmers
through certified seed programs, extension efforts, and other mechanisms (Duvick,
1983). Rather than breed for local conditions, breeders aimed for broad adaptability

of high-yielding fertilizer-responsive varieties in irrigated, fertile zones. Feedback
from farmers in on-farm trials rarely provided information on the suitability of
selection for specific locations. The seeds were delivered to farmers largely through
the patron–client extension model which focused on the individual farm enterprise,
not the community or social groupings of farmers (Duvick, 1986).
The “success” of the Green Revolution was double-edged. Signficant increases
in food production were achieved in a short time, leading to the alleviation of food
shortages and famine in critical areas (Mellor and Paulino, 1986; Plucknett et al.,
1987). However, with this success (along with other forces such as urbanization,
out-migration, grazing) and as farmers responded to markets, growth, and develop-
ment programs by adopting a few high yielding varieties, many landraces were
abandoned. Concern over genetic erosion by national and international agencies has
led to the creation of a global network of ex situ gene banks and living collections
where landraces and wild materials are kept in short- and long-term seed storage
(Plucknett et al., 1987). Fewer resources, however, have been given to support in
situ conservation by native communities, and even less attention has been aimed at
preserving the knowledge of local peoples regarding plants, a critical legacy just as
vulnerable to erosion (Nazarea, 1998a).
This historical ecological–evolutionary trajectory of traditional in situ manage-
ment of landraces underscores the following points:
1. The often controversial proposal to maintain the dynamic evolutionary management

of landraces within traditional landscapes is based on the historical reality of
marginal farmers as folk curators;
2. Despite the tendency of modern agricultural science to separate the genetic
“resources” from the local knowledge base, both are essential components of in
situ maintenance of diversity and, by extension, a well-supplied ex situ system; and
3. Despite the loss of diversity in the “favored” environments, rich gene pools still
exist in many farming communities which survive along the margins of the world
economic order. These marginal rural populations are often seen — even by
conservationists — as a threat to biodiversity in protected areas and surrounding
buffer zones. Our approach is to see them as part of the solution.
STRATEGIES FOR IN SITU AGROBIODIVERSITY CONSERVATION

BY INDIGENOUS COMMUNITIES
Over the years, we have spent a great deal of time working with subsistence
farmers in Asia, Latin America, and the American South (Nazarea-Sandoval and
Rhoades, 1994). We have studied these “native curators” intensively as anthropolo-
gists, worked with farmers as members of interdisciplinary teams at International
Rice Research Institute (IRRI) and International Potato Center (CIP), and — more
recently — as anthropologists attempting to document and revive landrace use in
the southern U.S. through support of traditional means of use and exchange of
heirloom varieties. Whether Andean farmers, Filipino rice cultivators and sweet
potato growers, or Appalachian gardeners, they share common characteristics. Uni-
versally, regions of rich biodiversity exist along the margins of their economic and

political worlds. Landrace cultivators are typically found in more remote mountains,
islands, rain forests, or desert agroecosystems which are momentarily insulated from
the dominant forces of the outside world economy (Dasmann, 1991). Communities
— and households within communities — with a propensity to maintain diverse
systems tend to be disenfranchised from the dominant order surrounding them. Even
the individuals who tend to be key native curators are marginal within their own
households. Thus, marginality at various scale levels is a key common designator
of landrace in situ curation.
Agroecologists, along with ethnoecologists who focus on the cognitive under-
pinnings of human–biological system interactions, have pioneered studies which
show that farmers pursue various strategies in using biodiversity as a way to meet
their basic physical, social, and spiritual needs (Hecht, 1987; Oldfield and Alcorn,
1991; Nazarea-Sandoval, 1995). This body of research points to fundamental dif-
ferences between informal and formal models of genetic resource/biodiversity man-
agement (Altieri, 1987; Altieri and Merrick, 1988). First, traditional producers/com-
munities use a different set of selection and evaluation criteria for germplasm
management than modern breeding or commercial seed programs. Second, their
methods of experimentation and testing are fundamentally different, although there
are some points of common interest. Third, the strategies which preserve biodiversity
are often embedded in community action which channels and encourages individual
households to act in such a way as to foster biodiversity.
Multidimensional Criteria for Selection and Maintenance of Landraces
Scientists find the tremendous diversity of landraces in marginal agroecosystems

useful and valuable in developing new and better varieties. To the practicing sub-
sistence farmer, however, it is strange — probably inconceivable — that one would
be so foolish as to risk this diversity with the narrow selection of just a few varieties
and species. In maintaining a wide range of varieties and species, traditional farmers
use multidimensional decision-making criteria which holistically involve ecology,
the complete food system from seed handling to consumption, and cultural aspects
such as culinary qualities, ritual, and cosmology. This complex decision-making
process may often be poorly understood by the formal scientific sector which tends
to be largely market oriented.
Farmers opt for an adaptive strategy of using biodiversity in such a way that it
spreads production risk and labor scheduling across the landscape. In the Cusco
Valley of Peru, for example, we found farmers who plant up to 50 different varieties
as well as several species of potatoes at different time intervals in 20 to 30 scattered
fields characterized by different altitude, soil types, and orientations to the sun. This
principle of diversity to spread risk is simply an Andean version of “don’t put all
your eggs in one basket.” This dispersion pattern reduces the risk that one disease
outbreak or an unpredicted frost will devastate an entire crop. Simultaneously, by
using different varieties a continuous flow of production through time and space can
be realized so that different markets, household needs, or labor supplies can be
accommodated. Interspecific and intraspecific variation is also used for agronomic
control of weeds and pests, microclimatic variation through shading, as well as a

buffer against climatic and pest damage. Andean potato farmers’ strategies are based
on a long-term, detailed knowledge of specific plant–environment interaction. Any
variety is tested against several seasons of variable frosts and rainfall as well as
performance in different soils.
In the Philippines, market forces are as salient as ecological factors in farmers’
decision-making frameworks, and the cognizance of instability and unpredictability
of both leads to constant experimentation, information and germplasm exchange,
and hedging. A sweet potato farmer in Bukidnon resists the pull toward monoculture
because of his perception that environmental flux and economic trends are beyond
his control. According to him,
I ask for different planting materials from our neighbors but I don’t mix them up. I
plant at least five different varieties of sweet potatoes at any one time to experiment
from which ones I get the most benefit. At different seasons, we should plant different
varieties because we never know which ones would be most productive (Nazarea,
1998b).
Some rice farmers, integrated as they are to the market system and credit infrastruc-
ture, still plant their favored varieties in the middle of clumps or at the borders of
agriculture extension and credit-backed varieties, thus managing to have their credit,
and eat, too.
In localized agroecosystems, household production units are also direct con-
sumption units; thus, they have a vested interest in carefully linking production and
consumption in a way not found in commercial systems where different activities
are typically carried out by separate groups. In subsistence systems, the household
unit manages all stages of the food system, including seed selection, production,
storage, processing, and marketing. Even when there is a need for interhousehold
exchange of genetic material, the linkages are generally along kin-based and com-
munity networks. There are no “formal” seed certification systems and the people
who select cultivars are the same ones who grow, process, store, eat, and
exchange/sell them. When the consumption unit and the production unit are
coterminus, a more-refined and more-detailed set of criteria is used compared with
when these two functions are separated.
In the Andes, an interdisciplinary research team from the CIP discovered some
39 criteria that farmers consider in their evaluation of varieties (Prain et al., 1992).
This led to the conclusion that farmers do not seek an ideal variety. Instead, farmers
seek to manage an ideal range of varieties that address their food system requirements
related to cash and subsistence needs (Prain et al., 1992). These requirements were
highly local and specific to household needs. In one of the research sites, for example,
farmers would grow “improved” varieties for subsistence while in another village
farmers cultivated folk varieties for the marketplace (Bidegaray, 1988). These unex-
pected uses were tied to certain local realities which only the farmers fully appre-
ciated. In one case, there was a shortage of land and wage opportunities so they
used their land to produce high-yielding varieties for food, while in the other case,
a nearby market provided higher prices for the valued native varieties (Brush, 1992;
Prain et al., 1992).

Another aspect of diversity maintenance involves postproduction activities (stor-
age, seed selection, processing, and cooking). Women, who are often in charge of
these nonfield activities, handle materials in such as way as to increase aspects of
diversity further. Shapes and colors proliferate in landrace material since these are
used as perceptual signals for sorting and identification. Most published research on
potato selection makes reference to the significance of differences in color, shape,
texture, and taste. Selection for “storability” or “culinary quality” occurs in the hands
of women who are acknowledged by the men to have superior knowledge of the
crop. Andean farmers are connoisseurs of potatoes which they evaluate with a wide
range of cooking descriptors as well as taste labels such as “flouriness,” “stickiness,”
“wateriness,” and so on. Native potatoes are universally recognized as superior to
improved varieties in terms of culinary quality.
Among Philippine sweet potato farmers, characteristics such as cooking quality,
aesthetic appeal, storability, and propensity of mixing well with other cultivars are
valued as much as yield or disease resistance by households surviving in the marginal
zones. Morphological, gastronomic, life habit attributes, familiarity gradients, and
functional criteria were used in distinguishing and prioritizing among varieties, and
were far from being mutually exclusive. Interestingly, local criteria for evaluation
of sweet potato varieties tend to be fuzzy or to trail off into gray areas as to which
properties or traits are positive or preferred and which ones are negative or not
preferred. For example, people would say they prefer sweet varieties but bland ones
are good to eat with fish and are a good substitute for rice during lean times, or that
newer varieties are desirable because they produce bigger roots but older varieties
produce tastier though smaller roots. The result of this “fuzziness” is that it is
impossible to construct a hierarchy of sweetpotato varieties from the most desirable
to the least, and, as a consequence, people retain different varieties in their farms
and home gardens.
Another dimension of genetic resource diversity in traditional societies often
overlooked by scientists and planners from more “utilitarian” urban-dominated soci-
eties is the interconnectedness between plants and cosmology, that part of culture
which deals with perception, ritual, religion, and worldview. Given the intimacy of
daily contact between cultivators and their biological environment, especially plants
and animals, a cultural interplay is not uncommon during which the domesticates
are assigned significant symbolic roles in the lives of the people themselves (Zim-
merer, 1991). Therefore, plants are more than just food. Plants are also ascribed
gender, spiritual qualities, mystical powers, and important religious roles in the lives
of the people (Down to Earth, 1994). People of many cultures believe they originated
from certain sacred plants (e.g., Mayan creation story and maize). In the case of
southwestern Native American groups, the diversity of maize types (and colors)
reflects group relationship, ethnic origins, cardinal directions, and a reverence for
diversity (Ford, 1984; Sekaquaptewa and Black, 1986).
Evidence from many cultures around the world points to a playfulness and
appreciation of landrace diversity as expressed not only through color and shape,
but also reflected in complex folk taxonomies and cultural identity related to lan-
draces. In the Andes, certain potato varieties are valued more for their symbolic role
in gift exchange and honoring guests at ritualized meals than for any agronomic and

economic values. Brush (1977) reports that the most highly prized varieties are often
the most delicate and least productive (see also Carter and Mamani, 1982). One
study from Bolivia pointed to the importance of potato diversity to the cultural
identity of the Aymara (Johnsson, 1986). In some parts of the Andes, the most
prestigious meal one can serve is made up of native cultivars, especially of potato.
Although such beliefs are frequently disregarded by scientists as superfluous, the
ethnographic record shows that such beliefs play a major shaping role in creating
variability among cultures (Zimmerer, 1991).
Try as we might, as scientists, to coax the fan of strategies into a logical, universal
framework, none seems to provide greater exploratory power than the “framework”
of expediency — of hedging, making do, and muddling through. By this, we mean
the development and maintenance of plant genetic diversity in local agroecosystems
based on day-to-day pragmatic concerns and the natural inertia that preserves diver-
sity due to the existence of a multiplicity of local demands and preferences but
cannot be fully satisfied by any one “ideal” or “best” variety. The decision-making
process, in other words, is characterized by conflicting demands, complementation,
and compromise, resulting in behavioral outcomes that augur well for the mainte-
nance of a wide variety of cultivars.
Comparison of Scientific/Formal Approaches to Biodiversity

Maintenance
Contrasting the approaches of traditional farmers and scientists in methods of

varietal selection can clarify the reasons plant-breeding programs often fail to reach
farmers with new genetic materials (Berg et al., 1991). Since traditional farmers deal
with holistic systems and multiple selection criteria they do not normally think in
terms of formal dichotomies like “improved” vs. “local” varieties. Farmers select
varieties that perform well in certain areas (e.g., agronomic, yield, marketability,
culinary) important to the context of their localized food system. Although farmers
do not use the agronomists’ multiple replications side by side, the folk selection
process is far from haphazard. Like breeders, traditional farmers have a systematic
way of seeking and integrating materials into their living, working informal gene
banks. Farmers are fanatic seekers of new varieties, and they will eagerly seek
materials wherever they can be found (e.g., formal seed programs, neighbors, mar-
kets). Once a new variety is obtained, it is generally planted on a small scale in a
kitchen garden or in a single row along the margins of a regular field. If the variety
proves itself, farmers amplify their production as the amount of seed allows. The
variety is observed and evaluated for multiple qualities relevant to the local food
system (see Table 1). All the while, they continue to maintain their own “germplasm”
bank which is constantly being replenished and experimentally culled (Rhoades,
1989a).
Many farmers are avid experimenters by nature (Richards, 1985; Rhoades and
Bebbington, 1995). The “atmosphere of experimentation” which characterized the
neolithic farmer since the earlier stages of cultivation is one of the foundations upon
which agriculture advances (Braidwood, 1967), and farmers are as creatively
involved in this ongoing process as are scientists. A key difference, however, between

Table 1 Breeders’ and Farmers’ Cultivar Selection Methods
Breeders Farmers
1. Genetically uniform cultivars (pure lines, clones, Genetically diverse cultivars
hybrids)
2. Test under ideal conditions Real-world conditions
3. Yield and disease/climate tolerance Multiple criteria; fuzziness
4. Widely adapted; agroecological target zones Niche-specific (rain-fed, poor soils,
(flat, irrigated, fertile, homogenous, inputs) inaccessible, local inputs)
5. Formal structures; highly centralized; top-down Informal; kin/community based; gendered
6. Negative attitude toward G × E Positive attitude toward G × E
formal and informal cultivar selection is that breeders tend to narrow the genetic
alternatives in search of yield and disease or climatic resistance while marginal,
subsistence farmers tend to broaden their choices by seeking more diverse varieties
to fit their overall needs (Soleri and Smith, 1998; Nazarea-Sandoval and Rhoades,
1994). Indigenous cultivators do not design, perceive, or manage plots or zones in
isolation of surrounding areas. To the contrary, they manage for diversity along
continuous boundaries by pursuing opportunities creatively to mix genetic resources
and inputs to meet their household and community needs. Farmers use diverse criteria
in selection and adoption decision making which does not necessarily end up with
the intentional elimination of “less desirable” options. What is desirable or not
desirable to local farmers may be a matter of taste, a matter of timing, and sometimes
a matter of mood. In other words, they use fuzzy multiple criteria; if not, the diverse
cultivars would likely have disappeared long ago (Nazarea, 1996).
One of the reasons that small farmers in marginal environments have benefited
little from the yield and disease resistance achieved by formal breeding programs
is precisely because of the real-world interaction between genotype and environment
(G × E). Breeding programs typically assume agricultural scientists know better than
farmers the characteristics of a successful cultivar (Witcombe et al., 1996). Breeders
select under favorable growing conditions, and, if adoption does not occur, the cause
is frequently assumed to be ineffective extension or insufficient seed production
(Ceccarelli, 1995). Breeding for broad adaptation to agroecological zones requires
large-scale centralized seed production and distribution which in turn further pro-
motes genotypes that might be inferior to the landraces they are replacing under
stressful conditions. This formal approach contrasts with that of marginal farmers
who have traditionally relied on a strategy based on both intraspecific diversity (crop
mixes and landraces on the same farm) and where seed is produced either on the
farm or obtained from neighbors through community-based informal seed networks.
In bridging the gap between breeding programs and farmers in marginal areas,
breeders have begun to think innovatively about marginal farmers, experimental
designs, field plot techniques, and landraces (Maurya et al., 1988; Galt, 1989;
Ceccarelli, 1995). As a result, participatory breeding programs have begun to emerge
in which farmers are encouraged through support and partnership with scientists to
exchange knowledge and test, under farmer experimental conditions and designs,
cultivars early in the breeding-selection process (Prain et al., 1992; Joshi and Wit-
combe, 1996). These participatory programs have already generated varieties that

match farmers’ needs and increase production simultaneously (Maurya et al., 1988;
Sperling et al., 1993; de Boef et al., 1993; Sperling and Scheidegger, 1995; Witcombe
et al., 1996).
SOCIAL, ECONOMIC, AND POLITICAL DIMENSIONS OF

IN SITU CONSERVATION
The Social Context of Community-Based Biodiversity Management
While unusual innovativeness in biodiversity preservation finds manifestation in

experimental, individualistic farmers, ultimately the social context of local commu-
nities shapes in situ biodiversity maintenance. In communities that have not yet been
fully incorporated into commercial markets and still manage high levels of landrace
biodiversity, future protection of such genetic resources requires values compatible
with locally defined social and economic goals. Whether biodiversity is decreased
or enhanced in the future may depend on the degree of self-determination of local
communities to attain these locally defined goals. In many traditional, closed cor-
porate communities (i.e., membership determined by birth), intergenerational equity
or “bequeath value” is as important as it is in developed countries where heads of
farm enterprises expect their families to continue to operate the business well into
the future. Many indigenous communities with a firm sense of place are aware of
the value and role of land and diverse crop inventories to their cultural survival, and
communally strive to guard these resources. Andean communities, for instance,
carefully regulate, through annual village assemblies, the rotation of land parcels,
the use of communal pastures, and the complex of species and varieties planted. In
other cases, such as in the South American Amazon, Amerindian groups purposely
increase biological diversity in many locations through shifting cultivation which
involves systematically transplanting crops throughout the forest (Castilleja et al.,
1995). The landscape in this setting is in fact a cultural creation by the populations
who have over the centuries altered the natural landscape through clearing, burning,
planting, and other mechanisms of diversifying resource clusters for their use. A
similar effect is obtained around the Awa Ethnic Forest Reserve in Ecuador where
the Awa Indians have planted “forest belts” of gardens and fields as a territorial
signal to logging companies and other outsiders (Castilleja et al., 1995).
Indigenous peoples value their community resources and typically practice col-
lective decision making at a much higher intensity than found in open, Western
societies. To maintain and exploit communal resources requires group values, ded-
ication, and willingness to follow village leadership faithfully (Rastogi and Pant,
1996). In most cases, regulation is enforced and punishment meted out by the
community itself. Leadership often rotates among households so that all will have
a degree of responsibility through time. Tribes of Arunachal Pradesh in India, for
example, have a self-governing system wherein the tribal council is responsible for
societal decision making (Rastogi and Pant, 1996). Their councils (variously known
as buliangs of the Apatanis, the kegangs of the Adis, the nyels of the Nishings) are

informal in nature and led by elders who command a great deal of respect. Any
conflicts, such as over boundaries or in sharing of resources, are handled in the
village courts through mediation. The main threat to these indigenous local institu-
tions has been the national and state governments which have declared control of
much of the ancestral lands by overpowering and disbanding local institutions
(Fisher, 1995). In Nepal, for instance, when the government seized all community
forest lands in the 1950s and superimposed outside government agencies to manage
the forest, widespread deforestation and land exploitation resulted (Ives and Mes-
serli, 1989). Even local people who formerly protected their lands participated in
the exploitation. Only in the 1990s, after a great deal of destruction, has the gov-
ernment tried to reverse itself and return the forests back to local community-based
management (Griffin et al., 1988; Gilmour and Nurse, 1991).
The power of communities in the social creation and maintenance of biodiversity
can be clearly seen through their role in seed or germplasm exchange of cultivated
crops through carefully organized fairs or market days. For centuries, people from
the native communities have congregated in spatially rotating markets on established
days during the year for the purpose of exchanging goods from different regions or
valleys. At Andean local markets and fairs, women with distinctive village apparel
sit at local markets behind sacks and baskets full of brightly colored beans, maize,
or potatoes. There are weekly fairs, which are tied to normal marketing of household
goods, and regional fairs held only once a year. The latter correspond to a religious
holiday and bring together communities from different zones and villages. These
festivals are held at the end of harvest so that a maximal diversity of crops, tech-
nologies, and knowledge is available for exchange prior to the commencement of
the next season. These regularly scheduled fairs, and the ritualistic manner in which
they are organized, in fact function for the systematic exchange of genetic materials
— a true manifestation of “conservation through use.” Similar markets are found
throughout the tropical world, such as certain mid-hill markets in the Nepal Hima-
laya which are renowned for the diversity of crops and the excellence of landrace
planting materials which arrive from distant valleys on a predictable schedule
(Rhoades, 1985).
The social locus of biodiversity management within households resides mainly
with women in households of marginal individuals. In our 1992 UPWARD project
in Bukidnon, Philippines, we compared two models for in situ conservation of
traditional root crops. One system was dominated by the traditional political hier-
archy, mainly comprising men, while the other was managed by an informal network
of migrant women called the “industrious mothers” (Nazarea, 1996). Through time,
the importance of the more informal, egalitarian approach of the women for biodi-
versity enhancement as opposed to the authoritarian formally organized male struc-
ture became apparent. The male organization could not sustain interest in the main-
tenance and enhancement of crop germplasm. In fact, genetic material in the male-
dominated garden would have perished had a female relative not rescued it by
transplanting surviving plants to her garden. The women’s garden, however, flour-
ished amid an atmosphere of lighthearted communal spirit from the beginning. All
members of the group, including children and men, joined in the process. A tracing
of the exchanges and information flows show that the women were connected in

layered vertical and interconnected horizontal relationships along which germplasm
and associated information were continually and reciprocally exchanged (see also
Prain and Piniero, 1994). One might surmise that the informal female network
replicates how germplasm has been created by traditional communities for centuries.
The germplasm is enhanced and conserved by different social “pathways” such as
between blood relations or ceremonial kinship networks or exchange between market
associates. Strong reliance on interpersonal and familial ties is important and a great
deal of germplasm changes hands between neighbors. Any new variety that is
acquired is soon spread, and redundancy considerably buffers the system from
genetic erosion. A well-entrenched cultural ethic of sharing, coupled with an interest
in seeking and soliciting materials to secure household needs, helps create and
maintain diversity.
Global Change and Plant Genetic Resources
The vast majority of marginalized indigenous communities in the world are still
unaware that during the past few years they have been thrust center stage in the
political debate over access, control, and ownership of the genetic resources found
within their traditional agroecosystems (Mooney, 1979; Fowler and Mooney, 1990;
Shand, 1991; Brush, 1993). Most are unaware of the concerted efforts of trade
liberalization and political policies aimed at their political and economical integra-
tion into the global “village” (Hall, 1991). However, this situation is rapidly chang-
ing. Due to a combination of rising awareness of the possibility of economic value
of locally controlled genetic materials as well as increasing legal and political debate
over the same resource on a global scale, marginalized farming communities are no
longer content to be partners voluntarily assisting plant collectors or ethnobotanists
in finding and identifying useful genetic materials in their territories. This newly
found awareness for traditional peoples further complicates an already entangled
global dynamic in which many stakeholders — private companies, national and
international breeding programs, activists, and academics — must now contend with
the role of indigenous communities that goes beyond their previous uncompensated
provisioning and protection of the germplasm in their traditional agroecosystems
(Rhoades and Nazarea, 1996). Indigenous communities can have crucial roles in
biodiversity protection if their ancestral territories and knowledge are legally recog-
nized, if they are provided effective control over the resources — including their
own knowledge — within their environments, and if they are not hampered by
external extractive policies which interfere with traditional life in a destructive
manner (Cunningham, 1991; Martin et al., 1996).
The modern thrust in development to empower local communities has been
generated, ironically, by a process called globalization. These changes are also
related to the new regional and supranational monetary and financial arrangements
(e.g., World Trade Organisation, North American Free Trade Agreement, etc.) which
break down trade barriers, open new markets, and bring democracy to formerly
highly authoritarian areas (Hall, 1991). One result of the global liberalization move-
ment is that, increasingly, local communities, even in formerly highly centralized
countries like China, are given more and more freedom to manage their local affairs.

These new arrangements signal possible new alliances between conservationists and
the international movement for local rights (e.g., human rights, community empow-
erment, self-determination). This often means that global environmental organiza-
tions endorse local community rights while the community adopts the slogans of
international environmental protection (even to the point of writing grant proposals
for funding).
In recent years, the policy recommendations and funding initiative following the
publication of Our Common Future have stimulated agricultural and development
agencies to show more interest in developing marginal areas of the developing world
(World Commission on Environment and Development, 1987). In many of these
hard-to-reach regions, a Green Revolution has not occurred, much to the frustration
of central governments, international agricultural research centers, and input suppli-
ers. Since such regions are not well integrated into national economies, they also
tend to be areas where political resistance, civil unrest, and narcotic trafficking or
illicit trade are common. They are also rich in many natural resources such as
economically significant flora (especially medicinals and timber), minerals, and
water. Despite their “invisibility,” the sheer numbers of such marginalized people
are not insignificant. Perhaps over 1 billion people live in the more marginal diver-
sified ecosystems that are somewhat insulated from “roadside” development —
mountain, uplands, arid and semiarid deserts, and tropical lowlands — where most
of the remaining world biodiversity is found.
Much of the development thinking about these marginal areas has been guided
by a “blame the victim” mentality; that is, too many unplanned children lead to
more poverty which in turn creates more land degradation which leads the vicious
spiral inevitably downward (Rhoades and Harwood, 1992). One widely promoted
solution to such land degradation, including loss of biodiversity, is to transform or
modernize such zones either through the transfer of technology or by enhancing
income through creation of enterprises which would allow the populace to market
their local resources on the outside. The farming system research and extension
movement was an effort in this direction, as is the more-recent participatory sus-
tainable agriculture and natural resource management model (Rhoades, 1989b).
Despite an emphasis on the value of indigenous systems in these well-intended
approaches, the prevailing belief among most agricultural scientists is that improved
agricultural systems should replace those traditional systems which are not capable
of producing sufficient food and income. Therefore, a great deal still hinges on the
development of external inputs and technologies (especially high-yielding varieties),
commercial markets, and road building.
The neoclassical economic approach to transforming agricultural communities
outlined in the previous paragraph has been questioned from the perspective of
agroecology and ethnoecology. Dove (1996), in discussing the granting of intellec-
tual property rights to farmers, even goes so far as to argue that well-meaning
projects which aim to help marginal producers maintain diversity actually serve to
undermine the diversity to be preserved. By pulling the marginal areas into the
mainstream, one destroys the context which fostered diversity in the first place,
that is, marginality. While other agro- and ethnoecologists may adopt a more

moderate position than Dove’s, many question the neo-Malthusian assumptions
about population–poverty–degradation by arguing instead that traditional commu-
nities possess creative and dynamic strategies for purposeful resistance to outside
influences (Nazarea-Sandoval, 1995; 1998a). Indigenous peoples are seen as cre-
ative actors who are becoming increasingly knowledgeable about global politics
and economics. This, in turn, has given them new external alliances which allow
them to add value to the resources they control. In other words, traditional com-
munities are no longer seen as “passive victims” in the global/political biodiversity
game (Colchester, 1994). In fact, many local communities are either closing their
doors to or negotiating with outside “bioprospectors” (Shyamsundar and Lanier,
1994). Recently, we witnessed in Ecuador the federation of Amazonian native
peoples halting all foreign collecting in response to an attempt by a U.S. researcher
to secure legal rights to a medicinal plant given by a shaman. Native peoples are
examining options by which they can gain economic leverage with both their
knowledge and the genetic resources found within their territories. This story is
only now unfolding in many previously isolated communities around the world
(Martin et al., 1996).
PATHWAYS TOWARD THE FUTURE
In the preceding sections we have outlined many of the problems and potenti-
alities of local in situ management and conservation of plant genetic resources. In
proposing positive future steps for effective conservation, we utilize a dichotomy
articulated by Janice Alcorn as a discussant at a symposium on “Local-global
(Dis)articulation of Plant Genetic Resources” (Rhoades and Nazarea, 1996). Alcorn
noted that in the rush to address critical issues of genetic resources, farmers’ rights
and intellectual property rights, poverty causal links, and other concerns, much more
emphasis has been placed on Conservation with a “big C” vs. conservation with a
“little c.” That is to say that more energy and debate are being directed toward the
macrolevel (e.g., international forums, world organization policies, national laws)
and much less is devoted to thinking about or working at the critical juncture, at the
microlevel. Alcorn concludes that we need more case studies, documented experi-
ence, and community models for working with farmer curators at the grassroots
level. Research on such case studies and models is now emerging at the global level
through a number of international programs (Martin et al., 1996).
Biodiversity will not be preserved in situ unless local communities see it in their
best interest to do so (Norgaard, 1988). However, this in itself is complicated. On
the one hand, marginality and poverty can drive communities to act in ways which
are not necessarily in accordance with the global agenda for biodiversity conservation
(Agenda 21 of the United Nations Conference on Environment and Development).
Many objectives of Agenda 21–inspired projects (nature reserves, endangered spe-
cies, clean air and water, natural resource management) may not be congruent with
the interests and priorities of local farmers (Reardon and Vosti, 1995). On the other
hand, subsidies, price supports, and other developed country approaches (e.g., such

as applied in soil conservation programs in the U.S.) will probably not work.
Solutions such as intellectual property rights, patents, and trade secrets will only
benefit local communities if the nation-state allows income from these legal arrange-
ments to reach local populations (Vogel, 1994).
While conservation of genetic variability has received national and global atten-
tion, funding continues to be inadequate to assure protection of ex situ collections
and many nations give low priority to genetic resource efforts. However, even less
recognition and funding support is given to preservation of the cultural knowledge
which underlies the genetic material (Nabhan et al., 1991). Interest in this activity
is relegated to a few token social scientists in international agencies or advocacy
groups such as local and international non-governmental organizations.
To address knowledge loss, Nazarea (1995; 1998a) developed an approach called
“memory banking,” based on the principle that effective germplasm conservation
should involve cultural dimensions along with the biological. She points to parallels
between memory banking and gene banking. While germplasm encodes genetic
information that has evolved as a response to selection pressures, cultural information
in the minds of local farmers who have had considerable experience in growing
these crops is likewise a repository of coded, time-tested adaptations to the environ-
ment. Moreover, genetic information embodied in folk varieties is threatened with
erosion because of pressure toward more intensive, monocultural production which
favors the adoption of newer, higher-yielding crop varieties in the same way that
cultural knowledge and practices associated with traditional varieties are in imminent
danger of being swamped by modern technologies (see also de Boef et al., 1993).
This loss of germplasm, and associated knowledge, has been referred to in a report
of a panel of the Board on Science and Technology for International Development
(National Research Council, 1992) as the “real tragedy of the commons; that is,
within a few decades traditional management systems in effect for thousands of
years become obsolete, replaced by systems of relentless exploitation of rural people
and rural countries.”
One of the most important issues for biodiversity conservation is how to give
farmers incentives to maintain diversity in light of these external pressures (Brush,
1993; Vogel, 1994). Although many marginal regions remain somewhat impervious
to the introduction of improved varieties as a result of inaccessible land and isolation
from markets, this same marginality leaves farmer curators normally politically
powerless and struggling with poverty. In such settings, the answer to in situ biodi-
versity preservation does not entirely lie with projects which focus on commercial
farms as private enterprise, but rather with the strength and interest of the traditional
community. In this chapter, we have already discussed such mechanisms as informal
curation, seed fairs, seed exchange gardens, and memory banking, and access and
control by local people of their native patrimony. If only a small percentage of
development funds could be earmarked to support individuals and the community’s
sense of pride in diversity, a significant amount of genetic variation might continue
to be nurtured in situ (Shulman, 1986).
Formulating effective guidelines or action plans for locally based conservation
is more easily written on paper than actually carried out in the field. We are fully
aware of the complications, trade-offs, and unanticipated effects, not to mention the

power of homogenizing forces of the global markets swirling around the remaining
pockets of agrobiodiversity on this earth. However, we cannot accept the conclusions
of the cynics who argue that diversity, once lost, cannot be restored or if the market
penetrates at all then the very comparative advantage of the marginal area for
diversity creation is gone. Brush (1993), Cleveland et al. (1994), and Nazarea (1996),
among others, show that diversity can indeed be reintroduced to regions and there
are many individual and project cases (e.g., Seed Savers Exchange, Native Seed
Search, Southern Seed Legacy) of both restoration and preservation.
In response to the apparent widespread loss of in situ genetic resources of
heirlooms (vegetables, fruit trees, and ornamentals) in the rapidly changing deep
south of the U.S., we established in 1996 a program called the Southern Seed Legacy.
What began as a limited project to test the memory-banking approach mentioned
above has taken on dimensions of a social movement itself. Farmers and gardeners
of diverse ethnic origins who were born in the earlier part of this century and whose
land is being enclosed by urban development still maintain diverse stocks of local
varieties of beans, melons, peas, okra, pecans, ornamentals, and dozens of other
disappearing varieties. Very little of this material is maintained within the region and
accessible to the local people in the U.S. Department of Agriculture germplasm
system. Likewise, other seed-saving organizations (for profit or not for profit) often
collect or solicit the southern seeds but maintain or market them from locations in
the northern U.S. or on the West Coast. Although the Southern Seed Legacy is a
young project, the enthusiasm for exchange and maintenance of the old varieties is
widespread and intense among both the old-timers and a new generation of small-
scale growers. A key to the success of the project so far has been the enthusiasm on
the part of farmer–gardener curators for documenting and passing along the crucial
link between cultural knowledge and preservation of the genetic material itself.
Regardless of whether in situ conservation is an indigenous event or one stim-
ulated by an outside-funded program, one principle must be preserved. That is, the
degree of independence, even irreverence, necessary for the persistence of diversity
is inversely proportional to the degree of integration into the market system and the
degree of capture by the political vortex. In situ conservation is a promising channel
of genetic conservation that is entirely compatible with ex situ gene conservation.
It already exists in the form of home gardens, polycultures, and traditional agroec-
osystems. The next step is to make this conservation strategy more systematic, more
sustainable, less risky, and to link it to imperatives beyond the local and even the
regional scale. Finally, if the system of access and rewards can be restructured, we
feel that local populations can use conventions that exist in the global marketplace
to their benefit — and for the benefit of humankind — instead of being subjected
to these conventions (Moran, 1998). Our argument is that marginal populations know
how to preserve biodiversity, have been doing it for centuries, and do not need a
great deal of outside direct assistance to get the job done. However, they cannot
continue to preserve the essential diversity solely for the sake of conservation if
there is no compensation for their labors or access and control over their ancestral
resources. These are the factors which need to be restructured. If not, we will “kill
the goose that laid the golden egg” and not even realize what we have done until it
is much too late.

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CHAPTER 13
Valuing Genetic Diversity:

Crop Plants and Agroecosystems
Douglas Gollin and Melinda Smale
CONTENTS
Introduction
Valuing Genetic Diversity for Agroecosystems: Concerns, Concepts,
and Cautions
Popular Concerns: Perception and Misperception
Conceptualizing the Sources of Economic Value
The Concept of Public Goods
A Caution: The Myth of Enormous Value
All Human Life Depends on Genetic Diversity
Some Species and Varieties Have High Value to Humans
Extinction Is Irreversible and, Hence, Infinitely Costly
Agricultural Genetic Diversity Protects Against Disastrous Disease
and Pest Outbreaks
Genetic Diversity for Agroecosystems: Distinctive Features
Approaches to Valuing Genetic Diversity for Agroecosystems
Farmer Decision Making and Farm-Level Diversity
Species Diversity
Varietal Diversity
Spatial and Temporal Diversity
Diversity, Productivity, and Stability: Hedonic Valuation and Related
Approaches
Genetic Diversity as a Public Good
Assessing the Value of Ex Situ Collections
Estimating Option Values

Implications for Conservation of Agricultural Diversity
Major Economic Questions Related to On-Farm Conservation
Conservation and Economic Development
Incentives for In Situ Conservation
On-Farm Crop Improvement as an Incentive
Research Impact
Property Rights and Conservation
Conclusions
Acknowledgment
References
INTRODUCTION
Researchers have long been aware of the importance of genetic diversity in

agroecosystems. At one level, diversity can provide ecosystem services to agriculture:
for example, beneficial insects and soil organisms can contribute to crop health and
reduce the need for agricultural chemicals.1 At another level, plant breeders and other
scientists rely on genetic resources for crop improvement and other technological
advances. Traditional plant breeding and newer biotechnologies make it possible to
incorporate desirable traits into crop varieties; this implies that crop landraces, wild
relatives, and other species have value as sources of desirable genetic traits.
To many agricultural scientists and other biologists, it seems self-evident that
genetic diversity has economic value and that genetically diverse ecosystems are
valuable resources. Many biologists take the view that, since all life depends on
genetic resources, their value must be infinite. The protection of all biodiversity (and
certainly of diversity in crop plants) should be a fundamental priority, in this view.
For example, Frankel et al. (1995) write that, “[T]he species serving humanity and
the communities safeguarding life and its diversity are of immense value. The highest
priority the human species can confer must go to their conservation. …” Other
authors voice similar views. Ehrenfeld (1988) argues specifically that economic
criteria are inadequate and inapplicable to thinking of the broader “value” of bio-
logical diversity:
Value is an intrinsic part of diversity; it does not depend on the properties of the
species in question, the uses to which particular species may or may not be put, or
their alleged role in the balance of global ecosystems. For biological diversity, value
is. Nothing more and nothing less. No cottage industry of expert evaluators is needed
to assess this kind of value. (p. 214; author’s emphasis)
Ehrenfeld goes on to argue that economists are fundamentally unable to account

for many of the most important aspects of value. One reason is the lack of adequate
biological knowledge about the functions of genes, species, and ecological commu-
1 See, for example, Tilman’s studies of the contributions that biodiversity makes to the level and stability
of biomass production per unit area (e.g., Tilman, 1997).

nities. A second reason that Ehrenfeld cites is the difficulty of putting values on
such intangible benefits as the satisfaction that people derive from the continued
existence of pristine environments. A third reason he gives is that the utilitarian
principle underlying economic valuations (the insistence on measuring net benefits
to humans) is inherently inadequate as a way to value the natural world. “The very
existence of diversity is its own warrant for survival,” Ehrenfeld argues. “As in law,
long-established existence confers a powerful right to a continued existence.”
Arguments such as Ehrenfeld’s have power, dignity, and deep ethical appeal.
Without taking issue with such viewpoints, this chapter will add to the “cottage
industry” of discussing the economic value of genetic resources. We focus specifi-
cally on the case of agroecosystems. Perhaps the justification for an economic
perspective is more readily apparent in the case of agroecosystems. Since agriculture
inherently involves the management of ecosystems for the benefit of humans, there
can be little reason to dispute the utilitarian premise of economic analysis. By the
same token, we have relatively little hesitation valuing genetic resources for agro-
ecosystems in terms of their current and potential use values.
Ehrenfeld’s first point, however, has substantial relevance for our work. Econo-
mists and agricultural scientists do not yet have much information about the extent
or potential value of genetic diversity for agroecosystems. There are few data yet
available about the degree of genetic variation within the Oryza genus for rice, for
example, or within the Triticum genus for wheat. Researchers are currently working
to map the rice genome, but it will be some time before the genetic bases for
resistance to major diseases and pests will be well understood. For more-complex
traits, such as yield potential, this understanding will require even longer.
In the meantime, important decisions must be made about the allocation of
resources for the protection and conservation of genetic diversity for agroecosystems.
How much is it worth to collect the remaining varieties of wheat from farms in
eastern Turkey? Should we build additional gene banks for ex situ storage of barley
varieties? Does the rationale for conserving wheat and rice varieties extend to spelt?
Does it extend to eggplant? To parsley? Is there a reason to encourage the cultivation
of “genetically diverse” plots of crop varieties or to encourage the conservation of
traditional varieties in farmers’ fields? Economists tend to approach such questions
with a particular concern for costs and trade-offs. Unlike some environmentalists,
economists generally believe that the costs of conserving genetic resources should
be viewed seriously, and that the benefits should be quantified to the greatest possible
extent (see, for example, Brown, 1990).2 Generally speaking, economists have been
more skeptical than biologists concerning the need to protect all forms of genetic
diversity. As Brown (1990) notes, “[I]f we can’t save all species, we need a ranking
based on one or more criteria, from which we select the highest ranked for conser-
vation.” This view — which would undoubtedly be considered heretical by most
biologists — is readily accepted by many economists.
2See also Evenson, R. E., Genetic resources: assessing economic value, manuscript, Yale University,
Department of Economics, New Haven, CT, 1993; also Wright, B. D., Agricultural genetic resource
policy: towards a research agenda, paper prepared for presentation at the Technical Consultation on
Economic and Policy Research for Genetic Resource Conservation and Use, International Food Policy
Research Institute, Washington, D.C., June 21–22, 1995.

To an economist, it is of central importance to consider the value of genetic
resources and genetically diverse agroecosystems (and even to assign dollar values,
where possible). Without such information, we are forced to make decisions on an
arbitrary basis. Even crude economic methods can offer valuable insights and can
help in setting priorities.
This chapter reports on recent conceptual and empirical work concerning the
valuation of genetic resources for agriculture, with particular attention to the role
of diversity within agroecosystems. The next section reviews current concerns over
diversity and summarizes economic theory and concepts pertaining to the valuation
of genetic resources. The third section reviews some of the empirical studies that
have sought to value genetic diversity for agriculture. The fourth section assesses
the implications of these empirical studies and other research for programs of
conservation of agricultural biodiversity. The last section offers some assessments
and concluding remarks.
VALUING GENETIC DIVERSITY FOR AGROECOSYSTEMS:

CONCERNS, CONCEPTS, AND CAUTIONS
Economists have long recognized the importance of biologically diverse farming

systems and have helped to draw attention to the multiple objectives of farm households.
A substantial literature in household economics and farming systems analysis has
analyzed farm-level decision making. Much of this literature has studied the portfolio
choice of farm households: in other words, the ways in which different farm and on-
farm activities are chosen to achieve multiple objectives, including high economic
returns and reductions in risk. Implicit in this literature is an interest in farmers’ efforts
to diversify production of agricultural crops through mixes of species and varieties. In
recent years, a number of studies have sought to assign a value to genetic diversity in
agriculture by showing how diversity is a private good; in other words, how individual
farmers benefit from incorporating genetic diversity into their farming systems.3 A
related question is the social benefit that is derived from diversity (for example, reduc-
tions in aggregate production variability that may be obtained if different farmers
choose different varieties, so that an aggregate level of diversity is maintained).
But genetic diversity has values that extend beyond the static function of diver-
sifying production and consumption choices. Increasingly, economists are focusing
their attention on the value of genetic diversity as an input into the production of
new crop technologies. Genetic resources are used to breed new crop varieties and
to improve agricultural technologies (e.g., pest control through Bacillus thuringien-
sis). Landraces and other crop varieties can be important future sources of disease
and pest resistance, and hence can be viewed as forms of insurance.4 For all these
reasons, the total economic value of genetic diversity extends beyond the short-run
private value that farmers attach to it.
3For example, farmers might benefit from growing varieties with different taste or consumption charac-
teristics, or from choosing varieties that are adapted to different microclimates or production locations.
4 Perrings (1995) articulates the view that biodiversity conservation should be viewed as a form of
insurance payment.

The following sections describe some of the concerns and directions of recent
economic research. We begin by considering the issues that have focused public
attention on genetic diversity in agriculture. We then ask how the concepts and
methods of resource economics can help to assess the value of genetic resources in
agroecosystems. Finally, we discuss the limitations of these methods.
Popular Concerns: Perception and Misperception
Much of the concern over genetic diversity for agroecosystems involves three
widespread (and related) perceptions. The first is the notion that modern agriculture
has caused genetic erosion, a term which encompasses the loss of traditional vari-
eties. The second is the idea that modern crop varieties and agroecosystems are
increasingly uniform, rendering crops extremely vulnerable to pests, diseases, and
other pathogens. The third is the view that genetic resources are scarce in agriculture,
i.e., that there is a problem with the adequacy of genetic resources for agriculture.
This last view is most commonly expressed in terms suggesting that modern agri-
culture is based on a precariously narrow stock of genetic diversity.
In all three cases, popular concerns are based largely on anecdotes. With regard
to the first concern, no causal relationship between the Green Revolution and genetic
erosion can be established for bread wheat, given the difficulties in measuring genetic
erosion on such a large scale and of demonstrating causality. The patterns of genetic
variation in farmers’ wheat fields have undoubtedly changed over the past 200 years
with increasing cultivation of varieties released by plant-breeding programs, but the
implications of these changes for the scarcity of useful genetic resources are unclear.
As expressed succinctly by Wood and Lenné (1997), the assumption that the spread
of modern varieties has been mainly responsible for an overall loss of traditional
varieties “goes beyond our knowledge of the facts of genetic erosion.” Historical
sources also demonstrate that most of the areas in which the Green Revolution has
had its greatest impact are high potential areas (not ancient centers of crop diversity)
which were targeted by local plant breeders and widely planted to the products of
their efforts since at least the early years of this century (Gill, 1978; Pray, 1983; van
der Eng, 1994).
With regard to the second concern, the evidence assembled in Smale (1997) and
Smale and McBride (1996) suggests that, in the major wheat-growing areas of the
developed and developing world, concentration among leading cultivars has tended
to decline as agricultural research and seed systems have matured. Semidwarf vari-
eties of bread wheat are in general more resistant to major pests and diseases, such
as the wheat rusts, than either traditional varieties or the taller varieties previously
released by breeding programs. They now incorporate broader and more-durable
types of resistance (Rajaram et al., 1996).
Finally, the stock of genetic diversity for agriculture remains relatively untapped.
There are many varieties (indeed many species) that have not yet been exploited.
Given the size of the genomes of agricultural crops and given the increasing feasi-
bility of incorporating genes from wild relatives or unrelated species through bio-
technology and other breeding techniques, it does not seem useful to think of genetic
combinations as determinate in number. In a recent study, Rasmussen and Phillips

(1997) show that genetic gains were made in barley based on a very narrow gene-
alogical base. They question the generally held belief that the variation on which
selection is based in elite gene pools is provided only by original ancestors, and
hypothesize that the contributions of newly generated variation and gene interactions
are underestimated. On all three counts, then, current concerns appear misplaced.
Conceptualizing the Sources of Economic Value
To point out the possible inaccuracies of popular concerns about the “loss” of
genetic material for agriculture is not to argue against the value of crop genetic
resources. From an economic perspective, genetic diversity has multiple sources of
value. A number of surveys have discussed these sources of value. (See, for example,
Brown, 1990; Pearce and Cervigni, 1994; Pearce and Moran, 1994; or Perrings et
al., 1995.5)
Perrings et al. (1995) suggest that it is useful to consider the total economic
value of genetic diversity as consisting of “use values” and “non-use values,” with
perhaps a broader valuation including noneconomic or “nonanthropocentric instru-
mental value.” Use values can be further disaggregated. One form of use value is
the direct benefits of genetic diversity to consumers and producers, such as the
increased satisfaction that comes from having a dozen apple varieties from which
to choose, or the increased productivity that arises through genetic crop improve-
ments. A second category of use value is the indirect benefits that people receive
from ecosystem functioning. Examples include the contribution of earthworms to
soil tilth, or of wetlands to flood control (e.g., Brown, 1990). A third type of use
value would include the future “option value” and “quasi-option value” of retaining
for the future the possibility of using a resource or of acquiring information about
that resource.
Use values are distinguished from non-use values (sometimes called “existence
values”), which reflect the pleasure that people derive from the sheer existence of
genetic diversity (without any regard for the usefulness of diversity). Thus, people
may derive some value simply from knowing that elephants exist or that rain forests
are being conserved. In considering genetic resources for agriculture, however, such
existence values are seldom of much importance. Relatively few people derive
satisfaction from the sheer existence of 80,000 rice varieties, but the use values are
substantial. By the same token, for other biological resources, existence values may
greatly outweigh use values. Few people derive much direct productive value from
the Bengal tiger, but many people value its continued existence.6
In most cases, then, it is use values that are relevant for agricultural genetic
diversity. More specifically, we will be interested in genetic diversity as it contributes
to expanded consumer choice and satisfaction and as it directly or indirectly enhances
5 See Swanson, T., The values of global biodiversity: the case of PGRFA, paper prepared for presentation
at the Technical Consultation on Economic and Policy Research for Genetic Resource Conservation and
Use, International Food Policy Research Institute, Washington, D.C., June 21–22, 1995.
6 It is worth noting, however, that economists have made great use of models in which consumers are
thought to have a “preference for variety,” such that they prefer to consume many different varieties of
a single product. Thus, for example, many people do appreciate (and are willing to pay for) diversity of
coffee varieties, apples, wine, etc.

producer profitability and security. These are private benefits that are obtained by
consumers and producers, and economic theory suggests that markets should do an
adequate job of meeting the demands for diversity that originate from these sources.
For example, consumers will be prepared to pay extra for exotic varieties of fruit
or (indirectly) for wheat varieties with special milling and baking characteristics.
Farmers will decide efficiently which varieties to cultivate and how to allocate land
and resources so as to achieve multiple production objectives.
But markets do not always work efficiently to provide genetic diversity. Genetic
diversity may generate benefits that cannot be captured by individual actors. In
such cases, markets may not provide diversity at adequate levels. Such cases are
discussed below.
The Concept of Public Goods
For most goods, market prices serve as measures of value. Prices reflect the
amount that people are willing to pay to purchase goods and services and the amount
that others are willing to accept in compensation for producing those commodities.
For some categories of goods, however, market prices are poor measures of
value. When consumers can benefit from a good without having to pay for it, the
market price will tend to understate the value of the good. Consider, for example,
an open-air concert in a public park. Many people can benefit from such a concert
without paying, and those who do pay (out of some sense of civic virtue perhaps)
will tend to contribute less than they might pay to hear the same concert in a concert
hall. Economists characterize goods such as this concert as “public goods.” Economic
theory predicts that freely operating markets will place too low a price on public
goods and will provide them at inefficiently low levels.7
Genetic diversity is a classic example of a public good. Although individuals
may benefit themselves from maintaining diverse farming systems, with multiple
species and varieties, their actions may also benefit others. For example, one farmer
may benefit from having her neighbor cultivating a diversified array of varieties,
reducing the attractiveness of their adjacent lands to certain kinds of pests or patho-
gens.8 But there is generally no mechanism through which diversity can be “pur-
chased” to ensure that it is provided in sufficient quantities.
Similarly, genetic resources themselves can be seen as a public good. Farmers
cultivate many traditional landraces of rice, wheat, and other crops. These landraces
represent resources that can be used by the whole world in the creation of new
varieties with desirable properties. But at present, there are no incentives beyond
7 The concept of public goods is discussed in any introductory economics textbook. The problem of
people sharing the benefits of public goods without paying for them is called the “free rider program.”
8 For example, suppose that there is a particular wheat variety that performs better than all other varieties
in a particular region (say, Minnesota). If every farmer in Minnesota grows this high-performance wheat
variety, it may increase the chance that a variety-specific pathogen will emerge. Each farmer then benefits
if some fraction of the farm population chooses to grow other varieties. But no farmer has an individual
incentive to grow a lower-performance variety. Moreover, there is no vehicle through which some farmers
can compensate others for growing the lower-performance varieties, even though everyone would expect
to benefit from such an arrangement. This is a common problem with public goods: the absence of a
market for a public good (genetic diversity, in this case) implies that the good will be provided at
inefficiently low levels.

their own private profit for farmers to cultivate these landraces. Although the global
community benefits from the conservation of genetic resources, there is at present
no market or other mechanism that would enable farmers to receive a share of the
global benefits. Without such an incentive, farmers will grow landraces only when
these varieties provide sufficiently high private benefits. The result may be ineffi-
ciently low rates of conservation of landraces.
A Caution: The Myth of Enormous Value
Markets may undervalue genetic resources and genetic diversity, but an equally
serious problem is that many people overvalue genetic resources. It is commonplace
for biologists and agricultural scientists to argue that genetic diversity has enormous
value. But such overvaluation of genetic diversity poses as much of a problem as
undervaluation by the market. If genetic diversity is indeed of enormous value, then
it becomes extremely difficult to establish priorities or to make trade-offs regarding
diversity. Without some estimates of relative value, we have no way to set priorities
for conservation or conversation. Some may argue that setting priorities is, in itself,
an unethical act of “picking winners” biologically. But as Swaney and Olson (1992)
write, “We are valuing biodiversity. We can choose to continue to undervalue [biodi-
versity], or we can change our valuations, but we cannot choose to not value it.”
Several arguments are advanced for the infinite value of genetic diversity. It is
worth looking briefly at these in turn. It is not possible in the space of a chapter to
characterize all of the arguments adequately, nor is it possible to refute them con-
vincingly. Instead, what follows is an attempt to explain the reasons economists are
skeptical about the alleged enormous value of genetic resources.
All Human Life Depends on Genetic Diversity
Some people argue that genetic diversity for agroecosystems is priceless, essen-
tially that it has infinite value. Without the genetic diversity of our crops, it is claimed,
humans would be unable to survive as a species; our agroecosystems depend on the
continued availability of a range of genetic resources. Although this is true, the high
total value of genetic resources should not be confused with the marginal or incre-
mental value of adding or subtracting one more species or gene. We are in no danger
of losing all the genetic diversity available to people, so the issue is best understood
as: How costly is it to lose some of the species or varieties now known to us (or,
equivalently, to protect some of the species or varieties now in danger)? The cost is
surely not infinite, but it is also not negligible.
Some Species and Varieties Have High Value to Humans
Lovejoy (1997) cites a number of instances in which genetic resources have

proved extremely valuable, from the directly useful genes found in penicillin and
perennial corn to the water filtration services provided by oysters in the Chesapeake.
There is no doubt that these examples are valid, but it is misleading to argue on the

basis of these examples that all genetic diversity is equally valuable (or even that
the average value is high). Many species have little potential for direct contribution
to human welfare, and many are sufficiently similar to other species that humans
might, realistically, suffer little from their loss. The valuable attributes of a particular
species might, upon search, be found to occur elsewhere or in other forms. Thus, it
is difficult to extrapolate at all from the “success stories” of biological resources
that currently have high values.
Extinction Is Irreversible and, Hence, Infinitely Costly
Biologists usually argue that the extinction of a species imposes losses on

humans. Two distinct effects are noted. First, an extinct species is “lost” for future
use, in the sense that its genetic materials cannot be put to utilitarian purposes. If a
species is extinct, we can never know whether or not it might have offered a cure
for cancer or — more prosaically — a gene that could be used in crop improvement.
Second, the loss of any species can perturb the delicate ecological balance of a
natural system. This in turn can cause damaging effects for humans.
In both cases, however, the costs of extinction can be overestimated if we do
not recognize the opportunities for people to find substitutes. As Simpson et al.
(1996) have pointed out, people can often find alternative sources of desirable genes.
There are arguably very few cases where a particular useful trait can be found only
in a single species or variety. More commonly, the compound occurs in several
species or varieties; or perhaps similar compounds are found in (related or unrelated)
species occupying similar ecological niches; or people can develop synthetic com-
pounds with the same attributes as the natural material; and so forth. The scope for
humans to substitute and adapt to extinction is remarkable. From the woolly mam-
moth to the passenger pigeon, humans have survived the loss of economically
important species without irreparable material losses.9
Agricultural Genetic Diversity Protects Against Disastrous Disease

and Pest Outbreaks
Within the agricultural sciences, a commonly cited justification for conserving

genetic diversity is the need to guard against potential outbreaks of diseases or pests.
Diversity gives farmers and scientists the resources with which to respond to emerg-
ing disease and pest problems. A related issue is the danger of genetic uniformity
in crops. Where cultivated varieties of a crop are closely related, it is suggested,
new pests and diseases can spread rapidly and with enormous destructive potential.
As evidence, several historical episodes are cited: the Irish potato famine, the South-
ern corn leaf blight epidemic in the U.S., and a handful of other well-documented
cases (e.g., National Research Council, 1972; Ryan, 1992).
9 The passenger pigeon is often cited as an example of an economically important species that was forced
into extinction by the failure of markets to reflect its increasing scarcity. Although it is certainly true that
market forces did not operate to encourage the conservation of passenger pigeons, it is equally true that
little long-term material harm to humans has resulted. This is not to deny that our lives are, in some
sense, poorer because passenger pigeons no longer fill the skies of North America.

Although genetic diversity undoubtedly is valuable for “maintenance breeding,”
it is probably not the case that the associated value of genetic resources is particularly
large. Typically, multiple sources of resistance can be tapped, and relatively small
arrays of genetic diversity would suffice to provide adequate resistance to many
diseases. Gollin et al.10 investigate this topic quantitatively, looking at the actual
distributions of resistance to diseases and pests in bread wheat. They conclude that
bread wheat landraces will be used for maintenance breeding on relatively rare
occasions, and they calculate as a consequence that these varieties have significant
but modest value.
Wright points out further that plant disease epidemics are rare and that their
costs are often overstated.11 Although the Irish potato famine was associated with a
blight, recent histories downplay the importance of blight as a cause of the famine.12
More recently, the Southern corn leaf blight epidemic barely caused a ripple.
Although this was due in part to the ready availability of resistant cytoplasm in
plant-breeding programs, it also reflects the inherent resilience of modern agriculture.
Trade, storage, crop diversification, and consumer substitution all serve to mitigate
the impacts of crop failures. Even in developing countries with imperfect markets,
farmers and consumers can rely on a variety of ex post consumption-smoothing
techniques to make up for the income losses associated with crop failures. (See, for
example, Rosenzweig, 1988; Rosenzweig and Stark, 1989; Udry, 1990; Alderman
and Paxson, 1992; Rosenzweig and Wolpin, 1993; Townsend, 1995.) As Sen (1981)
has argued in his seminal study of famines, crop failure does not correspond to
famine. Famine instead depends on a variety of other institutional and market failures
— often involving war, violence, or deliberate exploitation.
Genetic Diversity for Agroecosystems: Distinctive Features
The analysis presented in the preceding sections suggests that we should be

somewhat skeptical of exaggerated claims for the value of genetic diversity. But in
recent years, a sober literature has begun to seek theoretical and empirical measures
of the value of diversity. The dollar figures are not very precise, but they provide a
focus for discussion. The most common approach has been to value genetic resources
based on their “rareness” or their potential contribution to the development of new
or improved products. Among the more widely known efforts, Weitzman (1992;
1993) focuses on the measurement of diversity and on an application to the conser-
vation of crane species. Simpson et al. (1996) consider the likelihood that a new
species discovered in the rain forest will have value for pharmaceutical research.
10 See Gollin, D., Smale, M., and Skovmand, B., The empirical economics of ex situ conservation: a
search theoretic approach for the case of wheat, paper presented at the international conference: Building
the Basis for Economic Analysis of Genetic Resources in Crop Plants, sponsored by CIMMYT and
Stanford University, Palo Alto, CA, August 1997.
11 Wright, B. D., Agricultural genetic resource policy: towards a research agenda, paper prepared for
presentation at the Technical Consultation on Economic and Policy Research for Genetic Resource
Conservation and Use, International Food Policy Research Institute, Washington, D.C., June 21–22, 1995.
12 It is also interesting that few historians of the Irish potato famine consider the potato blight to have
been an important factor in the famine. For example, Mokyr (1983) does not include plant disease among
the seven “factors of importance” in explaining the Irish famine.

Similarly, Mendelsohn and Balick (1995) attempt to value undiscovered pharma-
ceuticals in tropical forests. This literature is hampered, however, by the paucity of
empirical data, which makes it difficult to move from models to empirically relevant
conclusions.
In contrast, the agricultural sector offers unique opportunities for describing the
value of genetic diversity. Agriculture has long used genetic resources to achieve
increases in productivity. Scientific plant breeding has been well developed in the
U.S. and Europe since the start of the 20th century, and organized systems for
collecting and conserving new genetic materials date back even further (Plucknett
et al., 1987). Government-sponsored systems of plant selection and varietal improve-
ment date back still further in Japan and China. Agriculture thus offers a logical
setting in which to examine the value of genetic resources. Moreover, data on plant
breeding and crop improvement are readily available for many crops, and much of
the research for the most important crops (wheat, rice, and maize) has taken place
in the public sector, so that data are accessible to researchers. By contrast, much
of the data on the utilization of genetic materials in, say, pharmaceutical research
are proprietary.
Studies of genetic diversity for agriculture must recognize some important dif-
ferences from the broader biodiversity literature. First, genetic diversity in agriculture
is in large measure a matter of intraspecies diversity, rather than interspecies diver-
sity. Second, agroecosystems are largely human managed; thus, the extent of diver-
sity is purposefully chosen, not the result of natural accident. Third, the conservation
of diversity (and, more generally, the conservation of genetic resources for agricul-
ture) intimately involves human communities. Agricultural diversity cannot be con-
served simply by setting aside tracts of uninhabited land; it necessarily involves
people. Rain forest biodiversity can be maintained (in principle) by conserving
habitat. But agricultural diversity can only be maintained in farmers’ fields as long
as incentives are appropriate.13
These distinctive features of agricultural genetic diversity have been recognized
in a growing literature concerned with valuing diversity. The next section of this
chapter summarizes some of the approaches and findings of this literature.
APPROACHES TO VALUING GENETIC DIVERSITY FOR

AGROECOSYSTEMS
The new economic literature on genetic diversity can be unraveled into four
distinct strands. One strand has sought to understand the choices of households as
they relate to on-farm diversity. A second strand of literature has focused on the
contributions of diversity to farm-level productivity. A third strand of research has
tried to measure the private costs of implementing a “socially optimal” level of
13For the conservation of wild crop relatives, of course, agricultural diversity does involve the protection
of uninhabited lands, as for wheats in the Near East and soybeans in China. Moreover, as W. Collins has
pointed out (personal communication, 1997), the protection of uninhabited lands raises problems for
agriculture, since protective measures often include prohibitions on plant collection (posing real problems
for agricultural scientists who seek to collect wild crop relatives for research purposes).

diversity. Finally, a fourth strand of literature has sought to identify the aggregate
value of genetic resources (or global diversity) to agricultural output. The four strands
of literature encompass both micro- and macrobehavior and address diversity at the
level of the farm, the region, and the globe. We consider these four strands in
sequence. The next section will consider the policy implications of these different
avenues of research.
Farmer Decision Making and Farm-Level Diversity
Agricultural genetic diversity is ultimately controlled by the decisions of farmers.

Farmers choose what crops and animal species to exploit and also select the mix of
varieties or breeds that they will use. Understanding farmer behavior is a prerequisite
for influencing agricultural diversity through policy, as well as for predicting the
impacts on diversity of external forces.
Species Diversity
The issue of species diversity at the farm level has long been viewed as a simple
question of production complementarities. Some agricultural commodities can be
produced together at lower cost than if they are produced separately. For example,
hogs can often be produced most efficiently on farms that also produce corn, soy-
beans, or other feed ingredients. Likewise, in many traditional Southeast Asian
agroecosystems, rice and fish can be produced together relatively efficiently. More-
over, some kinds of species diversity are economically efficient responses to uneven
income flows or uneven demand for labor through the year: livestock species may
provide off-season cash income; tree crops may offer a productive use for labor
outside of peak cropping seasons; and so on. Species diversification offers a shield
against production variation, price variation, and other forms of risk. Livestock and
tree crops may serve as a store of wealth, particularly in economies where financial
institutions are lacking. Species diversification can thus be viewed as an optimal
response to farm-level economic incentives.
Varietal Diversity
More of a puzzle has been varietal diversification: the tendency of farmers to

cultivate multiple varieties of a single crop. Varietal diversification is widespread.
Often, varietal diversification involves both “modern” and “traditional” varieties. A
long literature in agricultural economics has focused on farmers’ adoption of new
crop varieties (and on their continued use of old varieties).14 Initially, “partial
adoption” was seen as a puzzle: why would farmers persist in growing traditional
varieties alongside modern varieties? A simple decision-making model in which a
profit-maximizing firm chooses its technology from a set of available technologies
will give a unique choice, suggesting that a farmer should choose to grow only the
14This literature dates back to Griliches’s classic article (1957) on the diffusion of hybrid corn technology
in the U.S.

single variety with the highest net returns. But other explanations from economic
theory have been invoked to explain partial adoption. The most widely accepted
explanations involve (1) differentiation of varieties, such that different varieties are
in fact seen as different commodities, with partial adoption explained by some of
the same factors that explain species diversity; (2) farmers’ risk aversion, resulting
in portfolio diversification or disaster avoidance; (3) fixity or rationing of seed-
related production inputs such as fertilizer, soil type, or credit; and (4) learning
behavior or experimentation.
By examining the question in different terms, we can understand varietal “diver-
sity” as the counterpart of “partial adoption.” In the adoption literature, adoption is
viewed as a response to differences in yields and other characteristics associated
with modern and traditional varieties. In this literature, farmers choose the single
variety that gives the best set of characteristics. Instead, however, we can view each
variety (whether modern or traditional) as a bundle of characteristics or a multidi-
mensional vector of traits (Bellon, 1996). Farmers and their households derive utility
from these characteristics. Households seek to obtain desired levels of the charac-
teristics. To attain the desired levels, they may have to grow multiple varieties, so
that the allocation of crop area among varieties effectively weights their respective
characteristics.
In fully commercialized agroecosystems, market incentives may lead farmers to
choose bundles based on few characteristics (such as uniformity, a particular grain
quality, grain weight) — resulting in complete specialization among varieties. Vari-
etal diversity tends to be heightened when farmers produce both for their own
household consumption and for sale. In such cases, farmers may have multiple uses
for their crop and may seek to retain varieties with many different characteristics.
The “discarding” of varieties occurs when changes in the production environment
(or the tastes and preferences of the farm household) cause farmers to change the
weights that they assign to different characteristics. For example, if changes in the
value of women’s time encourage households to purchase commercial bread instead
of baking their own bread, farm families may be less inclined to grow wheat varieties
that are well suited to home baking and instead may grow more varieties with
commercially desirable milling characteristics. Varietal “loss” can also occur if many
characteristics of value to the farmer are bundled into fewer varieties, so that a new
variety satisfies many household objectives (Bellon, 1996).
Meng et al.15 suggest that partial adoption can be used to give a measure of the
value that farmers attach to genetic diversity. If farmers are continuing to cultivate
traditional varieties, even when modern varieties are available, it indicates that they
are willing to sacrifice the higher yield of modern varieties to gain some other
characteristics. The amount of yield forgone is, in this sense, a measure of farmers’
willingness to pay for retaining their traditional varieties. Meng et al. use data on
wheat cultivation from Turkey to consider a number of possible attributes or char-
acteristics that farmers may be “purchasing” when they choose to grow traditional
15
See Meng, E., Taylor, J. E., and Brush, S., Incentives for on-farm crop genetic diversity: evidence from
Turkey, paper presented at the symposium The Economics of Valuation and Conservation of Genetic
Resources for Agriculture, May 13–15, University of Rome Tor Vergata, 1996.

varieties: taste and baking properties, yield stability, and location-specific agronomic
features. The study concludes that farmers have a nontrivial willingness to pay for
the diversity that traditional varieties afford.
Spatial and Temporal Diversity
Farm-level diversity need not necessarily consist of multiple varieties or species

being utilized simultaneously on an individual farm. Diversity can equally be
achieved across space or over time. For example, if farmers grow one crop at a time
but alter varieties from year to year, and if different farmers are at different stages
in this cycle at a particular moment in time, then aggregate species or varietal
diversity may be high. Likewise, if farmers in different locations specialize in
different species or varieties, the aggregate diversity may be high even though no
individual farmer has a diversified farming system.
Spatial distributions are often affected by seed industries and delivery systems.
Spatial diversity can be represented by numbers of cultivars or percent distributions
by area planted to cultivars at a particular point in time. Changes in these counts or
distributions express the temporal diversity of varieties, or “diversity in time”
(Duvick, 1984). In mature commercial seed systems, temporal diversity, or high
rates of varietal turnover among farmers, substitute for spatial diversity (Plucknett
and Smith, 1986). Various measures of temporal diversity have been reviewed and
applied by Brennan and Byerlee (1991) and determinants of varietal replacement
have been treated in an analytical model by Heisey and Brennan (1991).
Diversity, Productivity, and Stability: Hedonic Valuation and

Related Approaches
The previous approaches are well suited to analyzing the impact of farm-level
incentives on diversity. But we can also ask how diversity contributes to productivity
(reduces productivity) at the farm level. Evenson (1996) notes that hedonic valuation
techniques may be useful in valuing genetic diversity and genetic resources as
producer goods. Hedonic valuation uses statistical techniques to assign value to the
characteristics of goods; it is the same approach, in effect, used by appraisers to
place a value on a house. The underlying principle is to observe how the value of
the final good changes depending on its characteristics. For example, appraisers
might observe how the sale price of a house depends on the roofing material; this
implicitly assigns a value to different types of roofing.
Similarly, it is possible to look at the productivity of rice in different localities
and to associate productivity levels with the characteristics of the breeding stock
used by plant breeders in that locality. Gollin and Evenson16 used this approach to
analyze the productivity of alternative categories of rice germplasm in India over
the period 1956 to 1983. The study sought to measure the relative contributions of
different types of genetic resources to varietal improvement and indirectly to pro-
ductivity change, using a two-stage estimation process that included clusters of
16
See Gollin, D. and Evenson, R. E., Genetic resources and rice varietal improvement in India, unpublished
manuscript, Yale University, Department of Economics, New Haven, CT, 1991.

genetic resource variables. Over the period 1972 to 1984, they estimated that varietal
change in rice contributed more than one third of realized productivity gains, while
public research and extension explained much of the remaining growth. Gollin and
Evenson found that the contribution of certain types of germplasm to rice produc-
tivity gains in India was very high. In particular, the early semidwarfing genes and
genetic resources associated with disease and pest resistance showed up as having
high value.
In the study by Hartell et al. (1997), the number of generations of plant breeding
and number of landraces in the genealogy of varieties were positively associated
with mean yield in rain-fed areas. These are indicators of genealogical complexity,
which can be viewed as a form of diversity. In irrigated areas, the concentration of
wheat area among fewer varieties was positively related to yield, while increasing
varietal age depressed yields.
Widawsky (1996) estimated the effects of varietal diversity on rice yields and
yield variability among townships in eastern China. He measured varietal diversity
using genealogical data and data on planted areas and concluded that diversity
reduced rice yield variability and only slightly reduced mean yields for the time
period under study.
In a sense, any study investigating the impact of plant breeding on yield is
analyzing the effects of genetic resources on productivity, broadly defined. Recent
studies of agricultural research impact, for example, have differentiated among
varieties based on their ancestry or the source of the germplasm. Bagnara et al.17
estimated the effects of local germplasm and international germplasm on the adapt-
ability, yield, grain quality, and yield stability of Italian durum wheats. Other exam-
ples include Byerlee and Traxler (1995), Pardey et al. (1996), and Brennan and Fox
(1995).
Genetic Diversity as a Public Good
As noted above, genetic diversity can be considered a public good, in the sense
that aggregate diversity may inhibit the evolution of new disease and pest biotypes
and may lead to greater aggregate stability in production and prices. Individuals
have no incentive, however, to consider the “socially optimal” pattern of diversity
when they make their varietal selections. Instead, they choose the variety or portfolio
of varieties that is individually optimal. At a regional or national level, the aggregate
of these individual decisions results in a level of diversity that may differ from the
level that is socially optimal.
Heisey et al. (1997) considered the case of wheat cultivation in the Pakistani
Punjab, where wheat rusts are an important source of yield losses. The rusts are a
family of pathogens noted for evolving rapidly in response to selection pressures.
In particular, planting of large contiguous areas with cultivars carrying the same
genetic base of resistance speeds the evolution of new rust biotypes. In turn, the
17See Bagnara, D., Bagnara, G. L., and Santaniello, V., Role and value of international germplasm
collections in Italian durum wheat breeding programmes, paper prepared for the CEIS-Tor Vergata
Symposium on the Economics of Valuation and Conservation of Genetic Resources for Agriculture,
Rome, Italy, Tor Vergata University, 13–15 May, 1996.

emergence of new rust biotypes can cause substantial losses to farmers and high
social costs if epidemics occur. From a social standpoint, then, it is desirable to
maintain some degree of diversity in the rust resistance genes incorporated in farm-
ers’ portfolio of varieties.
In practice, however, farmers do not choose to grow wheat cultivars with the
level of rust resistance that would be socially desirable. First, farmers choose to
grow high-yielding cultivars whether or not they are known to be susceptible to rust.
Second, farmers choose to grow high-yielding cultivars whether or not they have
the same basis of genetic resistance as those grown by other farmers. When many
farmers choose to grow the same higher-yielding cultivars, or when they grow
different higher-yielding cultivars with similar resistance genes, there is a lower level
of genetic diversity in farmers’ fields than the level that would most effectively
protect against the emergence and spread of new strains of rust.
Heisey et al. (1997) compared the portfolio of wheat varieties actually cultivated
by Pakistani farmers with an alternative portfolio and area distribution of cultivars
that maximized diversity, as measured by genealogical indicators. Switching from
the cultivars and areas actually planted to a more genetically diverse portfolio would
have generated yield losses worth tens of millions of dollars annually, even without
considering the costs of the policy interventions required to achieve it.
This research thus focuses attention on the supposed aggregate benefits from
diversity. Would the recommended portfolio actually perform enough better over
time to warrant the foregone yields? Does it matter how the recommended portfolio
is achieved? That is, does it matter whether every farmer grows each variety in the
recommended portfolio, or can farmers specialize entirely in the production of a
single variety? Are there other, more-effective ways of dealing with production
variability (e.g., through trade or stabilization policies) than with accepting reduc-
tions in yield? Are farmers individually better off growing lower-yielding (but more
stable) portfolios of varieties, or are they better off using ex post methods of income
smoothing to deal with production shocks? In other words, is on-farm diversity
optimal compared with alternative forms of individual or social insurance?
To address these questions, we clearly need some measurement of the benefits
of diversity as a public good. With good measures of these benefits, we could attempt
to assess the overall value of diversity. For now, this approach appears to offer an
innovative and intriguing framework for measuring the value of genetic diversity as
a public good. Continued research along these lines may prove fruitful.
Assessing the Value of Ex Situ Collections
Instead of considering the value of genetic diversity, per se, for agroecosystems,
we can think of valuing genetic resources themselves. What is the value of a collection
of genetic resources, which after all represent diversity in a latent form? To date, the
only empirical estimates of the value of a germplasm collection are in a study by
Evenson and Gollin (1997) that attempts to value the International Rice Germplasm
Collection (IRGC) at the International Rice Research Institute (IRRI) in the Philip-
pines. The general approach of the study is to associate the size of the IRGC with
international flows of germplasm and hence with increases in productivity.

Evenson and Gollin find that the size of IRGC influences the extent to which
national rice-breeding programs are willing to collaborate with IRRI experiments;
when countries think that IRRI is a source of valuable genetic material, they are more
likely to participate. Participation in the IRRI international experiments and germ-
plasm–sharing arrangements is in turn associated with increases in the rate at which
countries develop and release new varieties of rice. Countries that collaborate heavily
with IRRI, and that exchange genetic resources with IRRI and with other national
research programs, thus accelerate the process of technological change in rice.
Evenson and Gollin compute a value for additional accessions to IRGC based
on this relationship: they estimate that adding 1000 more cataloged accessions to
IRGC would generate 5.8 additional released “modern” varieties of rice (globally),
which would generate a stream of increased productivity worth $145 million annu-
ally (for a net present value of about $325 million).18
Several difficulties arise with these calculations. First, from a conceptual stand-
point, Evenson and Gollin are estimating only an “instrumental” value to IRGC
accessions; the value of a larger collection is simply that it stimulates greater
participation in the IRRI international collaborative programs. The study does not
attempt to value accessions based on the actual use of IRGC materials in the
development of new varieties. Moreover, the study assigns a value to accessions
based on the “average value” of a released modern variety, rather than on the marginal
value. This undoubtedly tends to overstate the value of IRGC.
Estimating Option Values
Other ways of valuing genetic diversity also warrant attention. As discussed

above, genetic diversity has an “option value,” which can be thought of as the value
that society might place on having the possibility of using genetic diversity at some
point in the future.19 The concept of option value is widely used in financial eco-
nomics. In financial markets, an option gives the purchaser the right to exercise a
particular choice at a future date. For example, an option might assign the purchaser
the right to buy a given quantity of wheat at a specified price at a time 3 months
from the present. Environmental economists have borrowed this idea as a useful
general framework for thinking about environmental goods: people may be willing
to pay a certain amount today to guarantee their right to make choices in the future.
There is no question that option values exist and are important: options are
widely bought and sold on financial markets. In principle, however, they do not
confer values distinct from productive values. Option values exist only insofar as
the goods or assets in question will have tangible value in the future. For genetic
resources, that value could be presumed to be future value in producing new crop
varieties or commercial products.
One interesting study that uses this concept was carried out by Brush et al.
(1992). This study provides evidence that Peruvian peasants maintain certain thresh-
olds of on-farm diversity even when the immediate advantages of switching to
18This assumes a 10% discount rate; at a 5% rate, the figure is $1.45 billion.
19Alternatively, we might consider the value that society would be willing to pay for information about
genetic materials at some point in the future.

improved varieties are large. These authors suggest that the cost of maintaining these
“emergency” stocks of traditional varieties represents a form of option value. This
is the amount that peasants are prepared to forego in order to maintain the option
of switching to other varieties at a later date.20
A related theme is embodied in the work of Gollin et al. (1997; see note 10),
who examine the actual distribution of economically important traits in the entire
population of wheat varieties. These distributions shape expectations about the value
of adding one more variety of wheat to the international ex situ collection. The value
of a variety, in this approach, is essentially an option value: it reflects the amount
that we expect the variety to be worth in terms of future yield losses averted,
independent of its current usefulness.
IMPLICATIONS FOR CONSERVATION OF AGRICULTURAL DIVERSITY
Ex situ and in situ strategies for genetic resource conservation are increasingly
viewed as complements rather than substitutes.21 Ex situ conservation is geared
toward a relatively small number of known plants. Ex situ conservation also “fixes”
the genetic material of the plant at the time that it enters the germplasm bank,
although genetic drift may occur over time. By contrast, in situ conservation can
be aimed at larger collections of species, some of which may not even be known.
Thus, protecting a rain forest is a form of in situ conservation. A virtue of in situ
conservation is that it allows adaptive, evolutionary processes to continue and
natural prebreeding processes to occur.22 Since the risk of extinction due to some
natural or anthropogenic process is greater in situ, ex situ collections serve an
insurance purpose.
Prospects for in situ conservation are likely to vary by species (Dempsey, 1996)
and are hotly debated for cultivated crops, on both biological and economic grounds.
For domesticated species or subspecies, in situ conservation implies farmer man-
agement of a diverse set of crop populations in the systems where the crops evolved
(Bellon et al., 1997). Conservation in this context refers more to maintenance of key
parameters in evolving systems rather than to conservation. Although the historical
role of farmers in shaping the evolution of crops and their diversity has long been
20 It is not clear, however, why individual farmers would not choose to “free ride,” in other words, to rely
on others to maintain the (unproductive) stocks of traditional varieties. Given that other farmers are
continuing to grow traditional varieties, an individual farmer has an interest in not growing them. Instead,
we might expect him or her to grow only modern varieties, secure in the knowledge that the varieties
will not disappear from existence. If all farmers behaved this way, of course, the traditional varieties
might in fact be threatened.
21 In common usage, in situ conservation involves the conservation of genetic materials in a natural habitat
or under cultivation. By contrast, ex situ conservation is the conservation of genetic materials in a

specialized storage facility (whether in an arboretum or botanical garden, or in a deep freeze, or in a
gene bank). In general, ex situ collections are intended to conserve genetic material outside of environ-
mental influences, whereas in situ collections are intended to conserve genetic material in its environ-
mental context.
22 Of course, in some cases, evolutionary processes may lead to the loss of genetic material and, hence,
may be undesirable from one perspective.

recognized, rigorous investigations of the complex socioeconomic and scientific
issues involved in such farmer-managed conservation efforts have only just begun.23
Major Economic Questions Related to On-Farm Conservation
Several economic questions of importance are raised by the prospect of farmer-

managed conservation of diversity. First, economic analysis of the costs and benefits
of conserving the intraspecies diversity of cultivated plants requires a different
conceptual approach than that proposed for in situ conservation of wild species. For
wild species, geographical areas may be isolated and contained in reserves, and
human populations paid or compensated for not harvesting, not grazing, or not
otherwise disturbing the plant populations.
The evolution of cultivated crops cannot be separated from that of humans,
however. Even in a center of origin and diversity, the crop genetic systems managed
by farmers are “open,” and their diversity relies on infusion of new genetic material
through seed exchange and varietal introductions, at least in the case of an outcross-
ing crop like maize in Mexico (Louette et al., 1997). Direct payments or other forms
of compensation to farmers for continuing to grow certain landraces have not yet,
in general, been proposed.
Conservation and Economic Development
The relationship of conservation to economic development is unclear. Many have

argued that on-farm conservation implies arrested development or, at least, cultural
stasis. Identifying conservation methods that would allow crop populations to evolve
without arresting economic development or penalizing farmers seems a laudable
goal (but it is by no means clear how to achieve it). Questions have been raised
about whether or not conservation can coexist with the integration of communities
into commercial markets.
Certainly the replacement of landraces with modern varieties is not an inevitable
process (Brush et al., 1992); farmers often continue to grow landraces even after
adopting modern varieties on part of their land. If farmers’ seed selection performs
a stabilizing function, then experimentation with introduced varieties, seed replace-
ments, and maintenance of landraces and their characteristics can coexist (Louette
and Smale, 1997).
Incentives for In Situ Conservation
Even if development is compatible with conservation under some circumstances,

what are farmers’ incentives to maintain the diversity they grow? From an institutional
23 At IRRI, the project “Safeguarding and Conservation of the Biodiversity of the Rice Genepool,
Component II: On-Farm Conservation”; in Mexico, the McKnight Foundation project “Conservation of
Genetic Diversity and Improvement of Crop Production in Mexico: A Farmer-Based Approach” and, at
CIMMYT, the project “Maize Diversity Management and Utilization: A Farmer–Scientist Collaborative
Approach”; in Turkey, the project “Ecology and Ethnobiology of Wheat Landrace Conservation in Central
Turkey”; a longitudinal study undertaken by the Institut National de la Recherche Agronomique (INRA)
in France; see other initiatives for Ethiopia and Andean crops described in Maxted et al. (1997).

perspective, Qualset et al. (1997) discuss various forms of direct and indirect incen-
tives for local conservation efforts. The research by Meng et al.24 is an attempt to
identify feasible policy incentives that support the management of genetic diversity
by Turkish farmers. In the Philippines, Bellon et al.25 were able to identify with
multidisciplinary methods a cluster of rice varieties that were both genetically diverse
and highly regarded by farmers in terms of consumption characteristics, disease
resistance, and other characteristics. These varieties were still cultivated in the rain-
fed system but had been “discarded” in the irrigated system. In this case, when both
the private and public benefits of growing the traditional varieties are high, the aim
of an on-farm conservation program would be to reduce the opportunity cost of
growing them, by, for example, shortening their growing cycle. This might be accom-
plished through rice breeding.
On-Farm Crop Improvement as an Incentive
In recent years, “participatory” plant breeding has been proposed as a means of

providing economic incentives for farmers to continue cultivating genetically desir-
able crop populations (see Eyzaguirre and Iwanaga, 1996). According to this point
of view, certain techniques used by professional plant breeders may assist farmers
to become more efficient in meeting their own seed selection criteria. Closer
farmer–breeder collaboration could promote yield increases or other improvements
in marginal environments where modern varieties have not been adopted for agro-
nomic, social, or economic reasons. Proponents of this approach argue that while
professional plant breeders have conventionally sought to develop fewer varieties
adapted to a wider range of environments, on-farm improvement can support the
maintenance of more-diverse, locally adapted plant populations — while including
farmers who had been “left out” of the development process.
Such approaches may only make sense in “marginal environments,” or, more
specifically, when the private benefit–cost ratio associated with growing traditional
varieties as compared with modern varieties is high. This would be true when farmers
do not have the choice of adopting an introduced cultivar that produces more benefits
(with respect to yield and/or any other combination of characteristics) than existing,
local cultivars — either because seed systems (formal or informal) don’t deliver
them or because their own materials perform better with respect to a range of
characteristics. Such a scenario appears to be the case in the Perales et al.26 study
of maize landraces in the Amecameca and Cuautla valleys of Mexico.
24 See Meng, E., Taylor, J. E., and Brush, S., Incentives for on-farm crop genetic diversity: evidence from
Turkey, paper presented at the symposium The Economics of Valuation and Conservation of Genetic
Resources for Agriculture, May 13–15, University of Rome Tor Vergata, 1996.
25 See Bellon, M. R., Pham, J. L., Sebastian, L. S., Francisco, S. R., Erasga, D., Sanchez, P., Calibo, M.,
Abrigo, G., and Quiloy, S., Farmers’ perceptions and variety selection: implications for on-farm conser-
vation of rice, paper presented at the International Workshop on Building the Basis for Economic Analysis
of Genetic Resources in Crop Plants, CIMMYT and Stanford University, Palo Alto, CA, August 17–19,
1997.
26 See Perales, H., Brush, S., and Taylor, E., Agronomic and economic competitiveness of landraces and
in situ conservation in the Amecameca and Cuautla valleys of Mexico, paper presented at the International
Workshop on Building the Basis for Economic Analysis of Genetic Resources in Crop Plants, CIMMYT
and Stanford University, Palo Alto, CA, August 17–19, 1997.

Since most of the approaches to on-farm crop improvement are labor-intensive,
the opportunity cost of labor is likely to play a decisive role in their attractiveness
to farmers. Labor is likely to be a factor in the attractiveness of selection techniques
for farmers. Rice et al. (1997) suggest that seed selection practices introduced for
maize competed for labor with coffee. Referring to a labor-intensive, on-farm selec-
tion program to improve seed quality and resistance to disease in beans, Sperling et
al. (1996) reported that the yield advantage of 14% was “not very convincing for
those who had to do the extra maintenance.” Zimmerer (1991) has argued that acute
labor shortages resulting from seasonal migration have undermined the management
of multiple crops and varieties in a traditionally complex cropping system in Peru.
In an Oaxacan community in Mexico, García-Barrios and García-Barrios (1990)
identify “diversity management” as first in a list of three agronomic characteristics
of pre-Hispanic maize systems that labor migration seriously threatens.
Research Impact
Even if a feasible innovation that is attractive to farmers can be identified, what

will be its research impact? Recommending strategies for on-farm maize improve-
ment or modified seed selection practices raises some very familiar issues related
to the adoption and diffusion of any agricultural technique, such as
1. Who adopts the practice and who does not?

2. If we observe farmers only for several seasons, can we conclude that they have
adopted the practice?
3. Is the practice adopted on both traditional and modern varieties?
4. Is the practice diffused from farmer to farmer or by other means?
5. Does adoption of the practice affect members of the farm household differentially?
6. Does the adoption of the practice generate any observable economic benefits, and
if so, for whom?
The benefits from any innovation depend on the way it diffuses among farmers
and the longevity of the innovation. Consider, however, the difficulty of assessing
the benefits of something like a new seed selection practice. To discern the benefits
of such a practice, we are concerned with the diffusion of new seed for the same
variety rather than new seed for a new variety. The conceptual frameworks and
analytical models social scientists use to analyze the factors that affect seed flows
(within varieties) among farmers are not nearly as well developed as those commonly
used to analyze the adoption of varieties. Further, as compared with the adoption of
variety, the adoption of a seed selection practice affects the characteristics of the
germplasm itself. To develop sensible approaches or models, we need answers to
basic questions, such as
1. How does seed flow among farmers, and to what extent is seed saved from
generation to generation or exchanged among farmers?
2. If seed is exchanged, what social “infrastructure” affects the direction and magni-
tude of its flows?

An appropriate analytical model must be based on the identification of the essential
social unit of seed conservation — which is not likely to be an individual farmer or
an individual household.
The research conducted by Aguirre,27 Louette et al. (1997), and Rice et al. (1997)
in Mexico demonstrates the high frequency of seed loss, seed replacement, and seed
introductions, both for seed of the same maize variety and for new varieties, including
both modern varieties and landraces. These studies, conducted in different regions
of Mexico with different research methods, reveal that only a subset of farmers
actually retains seed year after year (in spite of common stereotypes). Some farmers
replace the seed for their landraces deliberately, explaining that the seed is “tired”
or needs to be “rejuvenated.” Such practices have been cited in other literature, for
other crops (see Wood and Lenné, 1997). Other farmers mix seed from their own
and other sources, in search of “vigor.”
These findings are fundamental to understanding the genetic composition of a
crop and its diversity, and to understanding the diffusion and impact of techniques
to improve crops on-farm. They imply that the impact of on-farm improvement is
likely to be diffuse and difficult to observe, predict, and measure. To do so, we will
need to understand better the “social infrastructure” that shapes seed and information
flows, since in the diffusion of innovations of this type, the seed system is based
entirely on farmers and their interactions. The investigations of Ashby et al. (1995),
Sperling et al. (1996), and Almekinders et al. (1994) are examples of related research.
Property Rights and Conservation
An alternative approach to promoting in situ or on-farm conservation of diversity

is to invoke changes in property rights regimes that would allow farmers to claim
legal compensation when outsiders make use of landraces, traditional varieties, or
other genetic resources that have been developed by farmers. Farmers and indige-
nous people have, in many respects, created the array of genetic diversity for
agriculture that now exists. But at present, international property rights regimes
allow private sector firms to claim property rights over plant varieties created through
“scientific” breeding methods but make it impractical for traditional farmers to claim
comparable rights.
Several previous studies have documented the present system of property rights
for plant genetic resources and have surveyed the alternative forms of protection
that might be extended to indigenous peoples and to farmers in poor countries.
Specifically, Brush (1992), Gollin (1993a), Swanson (1994), and Walden (1995),
among others, offer overviews of current property rights regimes and analyses of
alternatives; Pray and Knudson (1994) investigate the impact of intellectual property
regime shifts on the composition of U.S. wheat production.28
27 Aguirre’s findings are reported in Analisis Regional de la Diversidad del Maiz en el Sureste de
Guanajuato, draft Ph.D. thesis, Universidad Nacional Autonoma de Mexico, Facultad del Ciencias,
Mexico, D.F., 1997.
28 See also a comprehensive review in Wright, B. D., Intellectural property and farmers’ rights, paper
presented at the symposium The Economics of Valuation and Conservation of Genetic Resources for
Agriculture, May 13–15, University of Rome Tor Vergata, 1996.

A number of changes in property rights regimes have raised the prospect that
farmers may soon be able to claim increased compensation for the use of traditional
varieties. Under pressure from developing countries, the 22nd session of the United
Nations Food and Agriculture Organization (FAO) arrived at an “International Under-
taking on Plant Genetic Resources” in 1983. This document declared that there is
a “universally accepted principle that plant genetic resources are a heritage of
mankind and consequently should be available without restriction.” This principle
was held to extend even “to newly developed varieties and special genetic stocks
(including elite and current breeders’ lines and mutants).” As Reid et al. (1993) note,
“[N]eedless to say, few developed countries with established seed industries sup-
ported the Undertaking.” In 1989, an FAO Conference rejected the previous formu-
lation and reached a compromise arrangement. Under the compromise, it was
accepted that property rights could legitimately be extended to breeders’ lines. At
the same time, the concept of “farmers’ rights” was recognized as a form of com-
munal rights “arising from the past, present and future contributions of farmers in
conserving, improving and making available plant genetic resources, particularly
those in the centers of origin/diversity” (cited in FAO, 1996).
Subsequently, the Convention on Biological Diversity, which took force at the
end of 1993, recognizes “the sovereign rights of States over their natural resources”
(cited in FAO, 1996). The convention also called for unspecified national measures
that would require signatories to share “in a fair and equitable way the results of
research and development, and the benefits arising from the commercial and other
utilization of genetic resources” with the countries providing the genetic resources.
The mechanisms by which such sharing would take place are not specified and are
taken to be worked out at the discretion of each pair of countries. It is unclear at
this point how far the convention actually extends; the U.S., for one, has not yet
ratified the convention. But it appears that the convention will work in the direction
of strengthening property rights protection for genetic resources (Gollin, 1993b).
The Convention on Biological Diversity specifically identified farmers’ rights as
an outstanding issue in need of resolution through the FAO Global System.29 Farm-
ers’ rights are defined in an FAO resolution as “rights arising from the past, present
and future contribution of farmers in conserving, improving and making available
plant genetic resources, particularly those in the centres of origin/diversity. These
rights are vested in the International Community, as trustees for present and future
generations of farmers, for the purpose of ensuring full benefits of farmers and
supporting the continuation of their contributions.”30 The FAO Commission on
Genetic Resources for Food and Agriculture is at present active in delineating the
scope of these rights, and it is likely that at some point a statement of farmers’ rights
will be proposed as a protocol for the Convention on Biological Diversity. At present,
it appears that if (when) farmers’ rights are given full international force, mechanisms
will be set up to provide compensation to farmers for the use of genetic materials
based on varieties that they have developed. There is currently much active debate
29 See Esquinas-Alcàzar, J., Farmers’ rights, paper presented at the symposium The Economics of
Valuation and Conservation of Genetic Resources for Agriculture, May 13–15, University of Rome Tor
Vergata, 1996.
30 Ibid.

about the mechanisms through which such compensation would be paid. Key issues
include the actual recipients of the compensation; the assessment of value; the
determination of genetic content; and the means by which “sharing of benefits” is
actually implemented.
Wright and Gollin31 suggest, however, that systems of farmers’ rights may in
fact provide little net compensation to farmers in developing countries. Many farmers
in developing countries are net “borrowers” of varieties from other parts of the world
and hence would lose rather than gain from a system of international compensation.
Moreover, the difficulties of implementing compensation systems are formidable.
For the purposes of on-farm conservation, it is unclear how monitoring would
proceed or how individual farmers (rather than governments or groups) would collect
compensation for the conservation or use of traditional varieties. Thus, on-farm
conservation may not be greatly enhanced by changes in international legal systems.
CONCLUSIONS
Genetic diversity has enormous value for agroecosystems, both as a resource

that can contribute to productivity change and as a source of “diversity services”
that aid farmers and consumers alike. Increasingly, economists have made advances
in understanding key issues involving genetic diversity for agroecosystems. These
include
Understanding the motivations and incentives that lead farmers to use diverse arrays
of species and varieties in their agroecosystems;
Recognizing the “services” that diversity offers within agroecosystems and the con-
tributions of diversity to increased productivity;
Accounting for the positive social benefits that farmers create through maintaining
diverse agroecosystems and through the conservation of landraces and traditional
varieties;
Understanding and estimating the prospective future value of genetic resources as a
source of desirable characteristics for varietal improvement;
Estimating the value of gene banks and germplasm collections; and
Identifying the key obstacles and policy issues relating to in situ conservation efforts.
In spite of these advances, many issues of importance remain unanswered. What

forms of diversity are most valuable? What are the priorities for research and for
conservation? How will changes in biological technology affect the importance of
diversity (both at the farm level and for researchers)? How best can individual
incentives be altered to reinforce individual incentives for conserving diversity?
31 See Wright, B. D., Intellectual property and farmers’ rights, paper presented at the symposium The
Economics of Valuation and Conservation of Genetic Resources for Agriculture, May 13–15, University
of Rome Tor Vergata, 1996; and Gollin, D., Conserving genetic resources for agriculture: local farmers,
international organizations, and intellectual property rights, paper presented to Globalization and Sus-
tainable Livelihood Systems Workshop, Institute for Social, Economic and Ecological Sustainability,
University of Minnesota, April 11–12, 1997.

Questions such as these will undoubtedly remain on the agenda for policy makers
in the years and decades ahead. Increasingly, however, economists are discovering
the methodologies and data needed to address these questions, and the prospects are
bright for future advances in understanding.
ACKNOWLEDGMENT
This chapter has grown out of the authors’ collaborations and conversations with
a number of colleagues over the years. These colleagues have shaped our thinking
and informed our views on genetic resources for agriculture. Among them are
Mauricio Bellon, Cheryl Doss, Bob Evenson, Paul Heisey, Dominique Louette,
Prabhu Pingali, Brian Wright, and members of the faculty workshop on biodiversity
at Williams College. Wanda Collins offered detailed and useful comments on an
earlier draft of this chapter.
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CHAPTER 14
Conserving and Using Crop Plant

Biodiversity in Agroecosystems
Wanda W. Collins and Geoffrey C. Hawtin
CONTENTS
Introduction
Sources of Genetic Diversity in Crops
The Need to Conserve Diversity
Approaches to Conservation
Ex Situ Conservation
In Situ Conservation
Integrated Conservation Strategies
Approaches to Breeding
Formal Systems
Informal Systems
Linking Conservation and Plant Breeding
Conclusions
References
INTRODUCTION
Biodiversity is the variability among living organisms from all sources, including
terrestrial, marine and other aquatic ecosystems, and the ecological complexes of
which they are a part. The interaction of various forms of biodiversity creates and
shapes the environment in which we live; it also creates and sustains the agroeco-
systems on which we depend for food and other basic needs. Diversity within an
ecosystem enables that ecosystem to survive and be productive and to produce an

enormous range of products and services. Agrobiodiversity is that component of
biodiversity that is important to agriculture and agroecosystems. It helps ensure
sustainability, stability, and productivity of production systems regardless of the level
of complexity of the ecosystem in which it occurs, and, in the final analysis, it
contributes to social welfare of the population through its contributions to poverty
alleviation and sustainable food security.
Diversity at the agroecosystem level contributes to greater food security, helps
increase employment opportunities, and increases local or national self-reliance by
allowing a variety of enterprises, based on products and services, to develop on a
national, regional, or community scale. A diversity of crop and animal species, at
the community, farm, or field level adds to social and economic stability through
reducing reliance on a single enterprise. Such diversity can also lead to a more
efficient use of natural resources, for example, through providing greater opportu-
nities for nutrient recycling (Carroll et al., 1990). Species diversity can also provide
a buffering effect against losses to diseases and pests or adverse weather conditions.
Diseases and insects that are major problems in large, single-crop species plantings
become less of a problem and cause less damage when additional crop species are
added to the system (Alexander and Bramel-Cox, 1991). Even at the field level,
diversity generated through planting crop mixtures can reduce losses to pests and
diseases. The net result of these types of utilization of crop diversity is resilience
and sustainability of agroecosystems.
Within any particular species, genetic diversity is the variation which is most
important. It is the variation that enables that species to adapt to new ecosystems
and environments through natural and/or human selection. Genetic diversity within
a cultivated crop species at the field or farm level helps diminish the risk of losses
through diseases or pests, and provides opportunities to exploit different features of
the microenvironment through, for example, the presence of diverse growth habits
and rooting patterns (Smith and Zobel, 1991). Such factors can contribute both to
greater stability and, in many circumstances, greater productivity. Both multilines
and variety mixtures have been used effectively for this purpose in grain crops
(Matson et al., 1997).
Modern agroecosystems often rely on crop species which are more uniform (i.e.,
less genetically diverse) than those in traditional agroecosystems. However, the
conservation of the existing genetic diversity of species is critical to the modern
farmer as well as the traditional farmer. Genetic diversity provides the reservoir of
genes for future crop improvement by farmers and professional plant breeders. The
ability to continue to rely on uniform, high-yielding crop species in modern agro-
ecosystems depends on the constant new identification and use of genes that are, or
have been, found in the genetically diverse crops of the traditional agroecosystems.
Similarly, useful genetic diversity can be found in wild relatives of crop species and
this diversity must also be conserved and appropriately used for improving crop
performance.
Thus, diversity is important to agriculture at all levels and in all agroecosystems.
While recognizing the importance to agriculture of diversity at all these levels, this
chapter will focus on genetic diversity within crops — the genetic resources that lie

at the heart of sustainable agricultural development and provide the basis for the
continued evolution and adaptation of crops.
SOURCES OF GENETIC DIVERSITY IN CROPS
Genetic variation within a crop gene pool can be found within and among
professionally bred varieties, landraces or farmers’ varieties, and nondomesticated
relatives. In addition, new genetic variation can be introduced through mutations
and the transfer of genes from different gene pools. Commercially released varieties
aim to combine genes for high productivity with those required to meet different
needs and environments. They contain a wealth of useful genes and gene combina-
tions and normally form the basis for further professional plant-breeding efforts.
Landraces and farmers’ varieties tend to be genetically heterogeneous and have
proved to be an excellent source of genes for, inter alia, adaptive characters and
disease and pest resistance. They are still widely grown, especially in marginal
environments where they may be more stable, and even more productive, than many
modern varieties. Landraces of many minor crop species are also still commonly
grown as, in general, they have received relatively little attention from plant breeders
and have been less subject to replacement by modern varieties.
Wild, nondomesticated relatives of crops frequently provide useful sources of
genes. For example, a wild rice, Oryza nivara, was used to introduce resistance to
grassy stunt virus in cultivated rice (Khush and Beachell, 1972). In Africa and India,
cassava (Manihot esculenta) yields increased up to 18 times after genes from wild
Brazilian cassava, conferring disease resistance, were incorporated into local vari-
eties (Prescott-Allen and Prescott-Allen, 1982). In the U.S., disease-resistant, wild
Asian species of sugarcane (Saccharum sp.) helped to save the U.S. sugar cane
industry from collapse (Prescott-Allen and Prescott-Allen, 1982). Many other cases
that have benefited agriculture in all parts of the world can be cited, as well.
In addition to these sources of genetic diversity, new DNA sequences can be
created or introduced into crop species. For example, mutations are a source of new
diversity and can be induced by chemical mutagens or ionizing radiation. And with
modern genetic engineering techniques, all organisms, at least in theory, can contain
potentially useful genes which could be transferred between crops and induced to
express themselves. These new genes then become integrated into the plant genome
and are passed from generation to generation.
THE NEED TO CONSERVE DIVERSITY
Conservation of genetic resources is essential, both to ensure that professional

breeders continue to have access to the genes and gene complexes needed for current
and future crop improvement and to enable farmers to continue to select and modify
their crops in response to changing environments and circumstances. Plant breeders
very effectively used genetic resources, coupled with higher levels of inputs, to meet

the demands of food shortages in the 1960s and 1970s. The result was the Green
Revolution. However, the lessons learned from that period have contributed to the
current emphasis on long-term sustainability of production systems and the protec-
tion of the natural resource base, while at the same time maintaining an adequate
and healthy food supply. This places new demands on plant breeders to continue to
increase productivity in socially and environmentally appropriate ways. Plant breed-
ers will respond to that demand; biodiversity is the means for achieving their goals.
Potentially valuable genes and gene combinations, which might at present be
unknown or undiscovered, can provide the means to fewer external inputs in crop
production systems, lower levels of environmentally toxic pesticides, and internal
resilience of agroecosystems. Conserving these needed genetic resources for future
use in the face of technical and political obstacles can be an enormous challenge.
The number of higher plant species is estimated to be between 300,000 and
500,000, of which approximately 250,000 have so far been identified (Wilson, 1988;
Heywood, 1995). Of these, about 30,000 are edible and an estimated 7000 have
been cultivated or collected by humans for food (Wilson, 1992). Despite this, the
crops that “feed the world,” by providing 95% of dietary energy or protein, number
only 30 (McNeely and Wachtel, 1988). Just three of them — rice, wheat, and maize
— account for almost 60% of the plant-derived calories in the human diet.
Much of the genetic diversity of major tropical food crops is now believed to
be reasonably secure in gene banks, especially of those species having orthodox
seeds such as rice, wheat, sorghum, and maize. For some crops, which require
conserving bulky vegetative organs or living tissues other than seeds, there are often
fewer accessions maintained. Cassava is a crop of world importance, but there are
only 23,000 accessions; and yam (Dioscorea spp.), an important staple crop in Africa,
is represented by only 11,500 accessions. There are still specific gaps in collections
even for crops which are well represented. For example, there are over 300,000
accessions of rice in storage, but there is still need for conservation of O. sativa
from Madagascar, Mozambique, South Asia, and Southeast Asia (FAO, 1996).
The situation is greatly different for most minor crops and those species that are
vegetatively propagated or produce seeds that cannot easily be stored for extended
periods at subzero temperatures. Crops such as taro (Colocasia esculenta), yams,
rice beans (Vigna umbellata), and breadfruit form part of the staple diet of millions
of the world poor, yet relatively little work has been done either to conserve or
improve them. In addition, there are many underutilized species of vegetables, fruits,
and other species, including nondomesticated plants, which contribute to nutrition
and dietary diversification in millions of tropical households. These species often
have been largely ignored because they are of little commercial value or because
they are at little risk of being lost. The result is that very little is known of the
diversity, distribution, and characteristics of such species and so conservation and
maintenance efforts are minimal.
Concern about the lack of knowledge and attention to conservation and enhance-
ment was expressed by many countries in the regional and subregional meetings
held in preparation for the Food and Agricultural Organization (FAO) International
Technical Conference on Plant Genetic Resources, held in Leipzig, Germany in June
1996. For example, representatives of countries in west and central Africa identified

a large number of underutilized species which are important to the livelihoods of
local populations. These included 7 species of cereals, 8 of legumes, 4 of roots and
tubers, 8 of oil crops, 31 of fruits and nuts, 17 vegetables and spices, 4 of beverages,
38 of medicinal plants, and 44 genera of forages.1
The situation for minor and underutilized species could be improved in the future,
as some gene banks have agreed to accept regional responsibility for long-term ex
situ storage of some minor crops. The National Bureau of Plant Genetic Resources
in India has, for example, accepted responsibility for rice bean, moth bean (V.
aconitifolia), okra (Abelmoschus esculentus), and amaranth (Amaranthus spp.), and
the Institute of Plant Breeding in the Philippines has accepted responsibility for
winged beans (FAO, 1996).
Because of the gaps in collections of both major and minor crops, the added
factor of genetic erosion increases the urgency to conserve diversity. The diversity
can be immense within the relatively small number of plant species which supply
most of the world energy and protein. For example, the International Rice Research
Institute gene bank contains about 80,000 accessions of rice. But much of the rice
diversity may already have been lost from farmers’ fields and may now only exist
in gene banks. By 1982, the rice variety IR36 was grown on about 11 million ha in
Asia and had replaced many local varieties (Plucknett et al., 1987). Although there
are varied reasons for the loss of genetic diversity, there is widespread agreement
that one major reason is replacement of local crop varieties by new cultivars. At
present, few quantitative data exist to define the extent and rate of genetic erosion
of crops and their wild relatives (Ceccarelli et al., 1992). Farmers in traditional
systems will routinely and intentionally discard components of local crop varieties
as a normal part of their management practices (Wood and Lenné, 1997).
The report State of the World’s Plant Genetic Resources (FAO, 1996) also lists
a number of other causes of such genetic erosion, including:
• Changes in agricultural systems and the abandonment of traditional crops in favor

of new ones;
• Overgrazing and excessive harvesting;
• Deforestation and land clearance, which is cited as being the most frequent cause
of genetic erosion in Africa;
• Adverse environmental conditions such as drought and flooding;
• The introduction of new pests and diseases;
• Population pressure and urbanization;
• War and civil strife; and
• Policy legislation (for example, until recently the cultivation of farm landraces was
discouraged in Europe).
Regardless of the reasons for disappearance of local crop varieties and their wild
relatives, the need to conserve that germplasm must be considered in conservation
strategies.
There are additional needs which must be recognized in considering the necessity
to conserve genetic resources, such as the high concentration of global collections
1 See http://www.icppgr.fao.org/srm/srm-syn/caf/E3.html.

for many export crops and commodities concentrated in a small number of countries.
For example, Zaire maintains over 80% of the global oil palm accessions. It is critical
that more attention be given to managing these collections, to safety duplication,
and to establishing new collections, especially in areas of diversity. In addition, some
tropical countries need to consolidate national collections to comply with the Con-
vention on Biological Diversity, which emphasizes in-country conservation of indig-
enous genetic resources, both in situ and ex situ. Many tropical and subtropical
countries report problems in maintaining collections because of the lack of suitable
drying facilities and unreliable electricity supplies (FAO, 1996). This situation is
particularly serious in coastal regions having 60 to 90% relative humidity. Further
constraints to conservation in the majority of developing countries are the lack of
adequate human and financial resources and infrastructure.
Overall, the efforts to conserve the most important world agrobiodiversity are
impressive and commendable. However, as noted above, there is still much to be
done and many questions to be answered about the most effective and efficient
conservation strategies. The need exists, as well, to ensure that agrobiodiversity is
not just conserved but also fully utilized to serve agroecosystem needs.
APPROACHES TO CONSERVATION
Conservation can be broadly considered in two ways: ex situ and in situ. Ex situ
conservation involves removing reproductive plant material from its natural setting
for maintenance in seed or tissue banks or plantations. Because of the finite nature
of any living plant material, ex situ conservation also requires regeneration of the
reproductive material at given storage conditions and at species-dependent intervals.
In situ conservation is accomplished by protecting plant material in the site in which
it naturally occurs. For most wild relatives this is in nature preserves or in wild
stands. For landraces, or traditional farmer varieties, it occurs in the fields in which
farmers grow those varieties (on-farm conservation) or in the communities in which
they are grown.
Ex Situ Conservation
Conservation in ex situ gene banks ensures that stored material is readily acces-
sible; can be well documented, characterized, and evaluated; and is relatively safe
from external threats. When material is stored in this way, plant evolution is effec-
tively frozen at the time of storage.
Of the main ex situ methods of conservation, the most common is the storage
of dried seeds in gene banks at low temperatures. For recalcitrant seeds, such as
those of many tropical perennial species, and vegetatively propagated germplasm,
such as Musa, cassava, or potatoes, other methods are needed. These include con-
servation as living collections in field gene banks or in vitro either as living plantlets,
as plant tissue on appropriate media, often under conditions of slow growth, or by
cryopreservation at very low temperatures, generally using liquid nitrogen. Genetic

resources are also conserved as frozen or freeze-dried pollen. Increasing use is being
made of isolated genomic DNA banks for the storage of germplasm (Adams, 1997)
and for most major crop species as cDNA inserts in microbial species such as yeast
and bacteria. DNA nucleotide sequence data for functional genes are a relatively
new mode of conservation of genetic resources. From these data, polynucleotides
can be synthesized in the laboratory.
For breeding programs, the screening of very large numbers of accessions for
specific traits can be expensive and time-consuming. Attention has been focused in
recent years on the development of core collections as a mechanism to facilitate
their use (Hodgkin et al., 1995). Core collections are collections that aim to represent
most of the diversity spectrum of the parent collection with a manageable number
of accessions, thus improving access to the whole collection. In setting up a core
collection, hierarchical approaches may be used, frequently with geographic origin
as one of the primary levels of discrimination. Specific adaptive traits (such as
maturity groups) can also be used to help stratify collections.
To be most useful to breeders, germplasm collections should be well docu-
mented. Accurate passport data, including site descriptions, are useful as a basis for
correlating origins with environmental parameters. Characterization data include
information on traits that are simply inherited and stably expressed in a wide range
of environments, such as major morphological features. These types of data assist
in discriminating among accessions and provide information on major adaptive
features (e.g., phenological characteristics). Evaluation data involve characters
important in crop production, such as yield and its components, resistance to dis-
eases and insect pests, flowering time, and plant height, and are perhaps the most
useful overall in the search for special adaptive traits, especially if originating in
diverse environments.
With respect to landraces, geographic origin and local knowledge can provide
very valuable leads to possible sources of genes. Farmers typically have a good
knowledge about the attributes of their varieties — e.g., phenology, reaction to
prevalent pests and diseases, and suitability for growing on the different soil types
found in the vicinity. Local knowledge is only rarely sought during collecting, and
greater efforts are needed to record such information (Guarino and Friis-Hansen,
1995). It has often been argued (IPGRI, 1993) that such information is under as
great or greater threat as the germplasm itself and data collection forms for plant
genetic resource collectors which provide for notes on ethnobotanical information
that should be obtained during collecting missions have been developed (Eyzagu-
irre, 1995).
With recent advances in computer science, not only are germplasm documenta-
tion systems becoming more powerful and user-friendly, but also data exchange and
the sharing of information among different systems are becoming easier. One exam-
ple of how the new technologies are being applied is the information system under
development by the International Agricultural Research Centers of the Consultative
Group on International Agricultural Research (CGIAR). Collectively, these centers
maintain over 500,000 germplasm accessions of most of the major world food and
forage crops. The information system is known as SINGER (System-wide Informa-

tion Network on Genetic Resources) and is available for international access through
the Internet.2
In Situ Conservation
In situ techniques allow the conservation of greater inter- and intraspecific genetic
diversity than is possible in ex situ facilities. They also permit continued evolution
and adaptation to take place, whether in the wild or on-farm where human selection
also plays a critical role. For some species, such as many tropical trees, it is the
only feasible method of conservation. Sustaining habitats indefinitely due to hazards
such as extreme weather conditions, pests, and diseases is a major concern for in
situ conservation. Difficulties in mapping, characterizing, evaluating, and accessing
genetic resources in situ are evident.
As with ex situ conservation, the method adopted depends on the nature of the
species. Traditional crop cultivars may be conserved on-farm, while undomesticated
relatives of food crops may require the setting aside of reserves. Agroforestry species,
and other plants which require little maintenance, can be conserved by developing
and maintaining sustainable harvesting practices and involving local communities,
while forest genetic resources are usually maintained in forest reserves and in areas
under specially designed management regimens.
One of the first steps for in situ conservation of target species or populations is
to determine their status in the area where they exist. It is also necessary to determine
the factors known to threaten the survival of the species and its vulnerability at
various stages of its life cycle. In the case of species threatened by extinction, the
minimum viable population size in the target area needs to be determined. This
concept implies that a population in a given habitat cannot persist if the number of
organisms is reduced below a certain threshold. The Species Survival Commission
Steering Committee of the World Conservation Union (IUCN) has recently devel-
oped new categories for threatened species based on population sizes, fragmentation,
and population viability analysis (IUCN, 1994). With the growing availability and
use of techniques for crossing plants which are distantly related and for transferring
genes from non related genera or even kingdoms, the search for useful genes has
been broadened. This has resulted in an increase in activities devoted to the collection
and maintenance of crop wild relatives (Ingram and Williams, 1987). This, in turn,
has led to a greater realization of the value of in situ techniques for ensuring the
conservation of a large range of potentially useful genes for future use in breeding.
Once considered primarily the domain of environmentalists and conservationists,
in situ conservation is now also becoming of increasing interest to those concerned
with crop improvement (Hodgkin, 1993). However, even though there is this growing
interest in the in situ conservation of genetic resources, most current in situ programs
target the preservation of ecosystems (often areas of outstanding natural beauty) or
particular species (generally endangered animals or plants) rather than the intraspe-
cific genetic diversity of plant species of potential interest for agriculture.
2 See http://www.cgiar.org/SINGER.

Options for in situ conservation range from nature reserves from which all human
intervention is excluded, through national parks in which economic activities with
a potential to disturb the natural ecosystems are carefully regulated, to the imple-
mentation of special management regimes in areas used primarily for agriculture
and forestry. The identification of specific areas in which a deliberate attempt is
made to increase and maintain intraspecific diversity of key species is another
approach (Krugman, 1984) which is being tried in Turkey and in Mexico. The Man
and Biosphere (MAB) program of the United Nations Economic, Social and Cultural
Organization is perhaps the largest coordinated global attempt to establish in situ
reserves, one of the objectives of which is the conservation of natural areas and the
genetic materials they contain. Under the MAB program, more than 250 biosphere
reserves have been established around the world.
As more attention is paid to in situ conservation, more innovative approaches
are developed. For example, locally based conservation (Qualset et al., 1997) seeks
to conserve biological entities at the farm, community, or regional level. Local issues
such as traditional and cultural behavior and knowledge play a large part in the
conservation effort and are thus conserved themselves.
Integrated Conservation Strategies
While plant breeders can readily access germplasm maintained in ex situ collec-
tions, it is far more difficult to do so in the case of material conserved in situ.
Nevertheless, the amount of inter- and intraspecific diversity that can be conserved
ex situ is a very small proportion of the total potentially useful variation. And, for
technical reasons, some domesticated and many wild species are very difficult to
conserve ex situ. Thus, to provide a comprehensive conservation program for any
particular species, strategies must include both ex situ and in situ approaches.
The comprehensive conservation of crop gene pools, which often comprise both
domesticated and wild forms, may require a combination of different methods, each
covering a different part of the gene pool, to enable the total to be conserved in the
most cost-effective and efficient way possible. Bretting and Duvick (1997) use the
terms static conservation and dynamic conservation (roughly comparable to ex situ
and in situ, respectively) to denote the purpose of the conservation programs rather
than the location. They recommend close collaboration between static conservation,
which serves to safeguard genetic resources outside the evolutionary context, and
dynamic conservation, which seeks to safeguard genetic resources in nature. In
dynamic conservation, the potential for evolution of the resources is conserved as
well as the cultural and agroecosystem properties that evolve along with them. The
two are not mutually exclusive but are seen to be integral parts of a continuum of
conservation.
The choice of appropriate strategies to protect and conserve the full range of
diversity in a crop species and its relatives depends on technical factors such as
reproductive biology and the nature of storage organs or propagules. It also depends
on the availability of human, financial, and institutional resources to sustain a course
of action once it is chosen. Such combinations of approaches are often referred to

as integrated conservation strategies and are based on the unique complementarity
of strengths and weaknesses between different approaches with respect to a single
crop (Hawtin, 1994).
APPROACHES TO BREEDING
Approaches to crop improvement generally fall into one or two broadly defined
systems:
• Formal systems, in which modern science is brought to bear on crop improvement

by institutions such as government plant-breeding stations, university departments,
and private breeding companies, with the aim of producing cultivars for wide, often
commercial, distribution to farmers.
• Informal systems, in which farmers and local communities, mainly in developing
countries, breed and select cultivars primarily to meet their own needs and circum-
stances.
Both systems coexist in many regions and each depends, to a greater or lesser extent,
on the other.
Formal Systems
Formal systems of crop improvement normally aim to produce high-yielding

cultivars that are broadly adapted across a wide range of agroecosystems. Special
attention is given to breeding cultivars with specific resistances to pests and diseases
and tolerances to abiotic stresses. Quality characteristics are generally determined
by the preferences of large consumer groups, often in importing countries, or by the
demands of processors. New cultivars must meet legal requirements for distinctness,
uniformity, and stability to be officially registered. Plant breeding is an expensive
process, and there is an ever-growing need to show financial returns on investment
in crop improvement from the public as well as private sectors. Under these circum-
stances, it is perhaps not surprising that the majority of modern cultivars are widely
grown or high-value crops and tend to have a relatively narrow genetic base.
In the formal sector, breeding for adaptation will continue to be concerned with
improving adaptability in existing environments, extending the areas in which indi-
vidual crops are grown, or seeking improved stability across a range of environments.
New variation will be sought and traditional cultivars and wild relatives will continue
to provide the necessary variation so long as their conservation is secured. Improved
knowledge of the distribution of diversity and of the effect of specific environmental
variables on that distribution will improve our capacity to locate desired characters.
Improved understanding of the significance and nature of coadapted gene complexes
will enable breeders to use adapted germplasm with much greater efficiency.
In formal systems, breeders are generally concerned with adaptation in one of
three ways:

• To develop cultivars that are better adapted to the agroecosystems in which the
target crop is currently produced;
• To develop cultivars that are adapted to new agroecosystems (geographic areas or
farming systems); or
• To develop cultivars that are competitive over large areas in the hope that they will
exhibit greater stability across seasons (by broadening adaptability).
The first of these is a more or less continuing part of most breeding programs and
includes breeding objectives such as the incorporation of improved frost resistance
in northern areas, better drought tolerance in arid lands, or disease and insect
resistance. The second objective has resulted in dramatic increases of area for a
number of crops such as sorghum in the U.S. (Maunder, 1992) or canola in Canada.3
The third has been a major objective of international breeding programs in the 1960s
and 1970s and resulted in such cultivars as the IR36 rice with its 75% coverage of
the Southeast Asian rice acreage. To achieve such objectives, use may be made of
genes having a large effect on specific, identified adaptive features, such as pheno-
logical characteristics; photoperiod response (e.g., day length in sorghum); tolerance
of extremes of temperature, soil moisture, or soil chemical factors; or resistance to
pests and diseases. However, a significant part is played by intensive directional
selection for characters under largely additive genetic control, such as flowering
time, maturation period, or by selection for broad adaptability as evidenced by low
genotype × environment interaction, as in the case of IR36 (Evans, 1993).
Informal Systems
In contrast to formal plant-breeding systems, farmer- and community-level crop

improvement efforts are more concerned with adapting cultivars to local conditions
and systems. In regions of high ecological diversity, a large number of microenvi-
ronments may have to be catered to, and farmer circumstances change over time.
In continuing to adapt local cultivars to meet their needs, farmer selection pressure
may have the effect of producing a series of small adaptive shifts, primarily within
the context of complex gene systems. Their high levels of heterogeneity compared
with modern commercial cultivars, comparative stability across seasons, location
specificity, and generalized, rather than highly specific, tolerances and resistances
often generally characterize farmer varieties or landraces.
Brush (1995) has shown that farmers in widely differing environments continue
to grow and use traditional cultivars on at least part of their land, even when modern
cultivars are available. Traditional cultivars (landraces) may meet their specific
requirements and may be better adapted to the environments in which they are grown.
There is growing evidence (Riley, 1996) that throughout the world farmers, partic-
ularly in traditional agricultural systems, actively seek to improve their crops through
exchange with other farmers and through selection within and among their landraces.
There have been reports of farmers making crosses, and gene flow between crops
3 See http://www.canola-council.org.

and their wild relatives can provide additional sources of variation especially in
centers of origin, although the extent to which this occurs has been questioned (Wood
and Lenné, 1993). Thus, landraces, or farmer cultivars, are not static, but represent
diverse and dynamic gene pools that, under the pressures of both farmer and natural
selection, evolve over time.
Smallholder farmers, particularly those in marginal environments, have to cope
with great environmental variability. They generally lack the economic and institu-
tional resources to transform their environments to meet the requirements of their
crops (Eyzaguirre and Iwanaga, 1995). As a result, they rely on within-landrace
variation or the inherent plasticity of their crops to ensure at least a minimum level
of production over the seasons. For many, meeting immediate family needs and
short- or medium-term survival are primary objectives. Strategies for achieving them
frequently depend on the maintenance and management of diversity within and
among crops and landraces in their farming systems. They must balance, for example,
straw vs. grain yields, high yield vs. the ability to yield under stress conditions, and
ease of hand-harvesting vs. optimum plant architecture for mechanical harvesting.
Under these circumstances, farmers’ strategies for managing their genetic resources
have resulted in an enormous diversity among and within their crops and cultivars.
LINKING CONSERVATION AND PLANT BREEDING
To serve the two broad crop improvement systems, the formal and the informal,
different approaches are needed in the conservation and supply of the diversity for
current and future genetic advance. As a generalization, ex situ methods were
developed primarily to meet the needs of the formal crop improvement system. The
importance of in situ conservation of crop wild relatives was recognized, but, until
recently, there were very few programs that specifically targeted wild relatives of
crops. Within the informal system a more holistic approach has been advocated and
activities aimed at supporting the conservation of traditional crop cultivars in situ
have been initiated. As discussed earlier, there is a great need and opportunity to
promote an integration of these approaches.
Conservation per se is rarely a conscious objective of farmers; however, in
applying selection pressures for particular traits, traditional farmers are generally
aware of the need to maintain high levels of “background” diversity. The ways in
which traditional farmers manage their genetic diversity have dual effects of con-
serving within the gene pool many of the adaptive features of their crops either
individually or as coadapted gene complexes, while at the same time allowing that
gene pool to evolve in other respects to meet new needs.
Plant breeding in the formal sector depends on four main sources of diversity:
wild relatives (and increasingly nonrelated taxa), mutation (often artificially
induced), landraces, and modern cultivars. Of these four, the variability in landraces,
or farmer varieties, is probably the most underexploited. Isozyme and molecular
analyses show that landraces and wild relatives remain the main reservoirs of genetic
diversity in crop gene pools (Miller and Tanksley, 1990). In addition, much of that
diversity may not be obvious even through traditional genetic manipulations, but

may be located and successfully used by employing molecular analyses (Tanksley
and McCouch, 1997). Genetic erosion of these reservoirs continues, in some cases
at alarming rates and for various reasons, and it is imperative that efforts to conserve
them be strengthened. Ex situ gene banks alone are not sufficient, and it is in the
best interests of both the formal system as well as the traditional farming commu-
nities themselves that the dynamic “cauldron” of genetic diversity be maintained in
situ, and be allowed to continue to evolve.
Formal breeding, for economic reasons, has tended to neglect crops of only
local importance and the needs of farmers in diverse marginal environments. How-
ever, for farmers in marginal areas, such crops may be the basis of their survival.
They have to depend on their own efforts and can rarely expect assistance from the
formal system. It is highly unlikely that formal breeding will ever fully meet their
diverse needs.
The need to produce more food for the ever-growing world population requires
increased productivity from all sectors. Both formal and informal systems of crop
improvement have a vital role to play; yet each tends to operate independently of
the other. While there is a flow of materials from the informal to the formal system,
its genetic diversity remains inadequately exploited by professional breeders. Ways
must be found to link these valuable sources of diversity more closely to professional
breeding efforts.
Qualset et al. (1997) suggest ways that both farmers and professional breeders
can link their plant-breeding efforts in breeding for “locally based conservation and
improvement” of crop plant genetic resources. In a joint breeding effort, farmers
would continue their tradition of improving their own material within the limits of
their resources. Formal breeding programs would contribute by introducing new
selection criteria and techniques that would improve performance but not change
the basic character of the crop. Single gene traits which could, for example, improve
disease resistance could also be introgressed and be made available to farmers in
the genetic background of the material they are already using. Sperling et al. (1993)
successfully involved farmers in Rwanda in breeding and selecting segregating
populations of beans.
Integration of formal and informal breeding efforts are increasing and will
stimulate the flow of genetic resources and information in both directions. Additional
ways in which the formal system can assist the informal include the following
(Hawtin et al., 1996):
• Strengthening links between farmers and gene banks to ensure adequate long-term
conservation of landraces, with systems to facilitate the restoration of landraces to
communities that have lost them;
• The provision by gene banks of adapted materials collected from one location to
farmers in other, environmentally similar locations for local testing and adaptation;
• The provision to local communities of segregating populations and other breeding
products derived from their own landraces;
• The introduction into local landraces of specific genes of relevance to local cir-
cumstances;
• The provision of a broad range of elite lines, within and from which farmers can
select according to their needs; and

• Training farmers in crop improvement techniques that are relevant to their own
circumstances.
As already mentioned, more participatory breeding approaches are being devel-

oped and tested in which formal sector breeders are working closely with farming
communities. Through a greater understanding of the needs and aspirations of such
communities, and through working in partnership, scientific expertise can best be
brought to bear on the problems faced by those farmers that have been the most
neglected up to now. Such approaches are expected not only to contribute to rural
community development, but may also help to ensure that the large gene reservoir
managed by the farmers continues to exist and to evolve, and remains a resource
for all crop improvement efforts in the future.
CONCLUSIONS
The contributions of agrobiodiversity to a variety of agroecosystem functions

and characteristics are well documented. The need to conserve genetic resources as
the fundamental source of agrobiodiversity is also well documented. Targeted and
efficient conservation strategies for the array of plant species that contribute to
agrobiodiversity will involve integrated approaches which use the best approach for
each species and in each system where diversity is found.
To use existing genetic diversity most effectively in an array of agroecosystems,
both formal and informal plant breeding systems are vital. Just as an integrated
approach to conservation will result in more efficiency in protecting genetic
resources, so will integrated approaches to plant breeding result in efficiency of
utilizing genetic resources and better meeting the various needs of agroecosystems
with respect to biodiversity.
Research and development activities which serve to link conservation and use
more closely will promote the realization of both objectives. The involvement of
farmers and farm communities, as holders of valuable genetic resources, in those
activities at early stages fosters the dual objectives by preserving and enhancing
traits which are often ignored by the formal sector, while at the same time providing
access to much-needed productivity increases. The appropriate, locally determined
balance between the two will lend stability and sustainability to both crop production
and crop genetic resource conservation. By blending the effective use of locally
adapted landraces and infusing genes for specifically needed production traits, the
valuable, farmer-held genetic material will continue to be conserved, on mutually
agreed terms, for potential use by formal sector breeders.

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CHAPTER 15
Implementing the Global Strategy for

the Management of Farm Animal
Genetic Resources
Keith Hammond and Helen W. Leitch
CONTENTS
Introduction
Evolution of the Global Strategy
Principles of AnGR and the Rationale of the Strategy
Imperatives for Action
The Global Strategy
From Strategy to a Global Initiative
Benefits of the Global Strategy
Strengths and Innovative Features of the Global Strategy
References
INTRODUCTION
Domestic animals meet 30 to 40% of human needs for food and agriculture.
Over 1.96 billion people derive some livelihood from farm animals, and, for 12%
of the world population, domestic animals are the only assurance of food security.
By the year 2030, world food production must increase by more than 75% in order
simply to maintain current levels of food availability. Animal production must
increase by at least this amount.
Domestic animal genetic resources (AnGR) underpin food security, yet irreplace-
able resources are disappearing at an alarming rate: about 30% of the estimated

5000 breeds, comprising some 40 species of livestock, are at risk of extinction.
About 80% of these indigenous breeds at risk occur only in developing countries
and few-to-no conservation programs are in place. The fact that agriculture produc-
tion depends on so few AnGR magnifies the importance of genetic diversity repre-
sented by those remaining — diversity that provides farmers with the raw material
to develop livestock to be more productive; more able to resist diseases or better
adapted to adverse conditions defined by available feeds, climate, and many other
stresses; and more able to respond to the changing needs of modern communities.
AnGR is a key sector of agrobiodiversity. Biodiversity enables the sustainable
development of agriculture. By definition, sustainability is production environment
specific, and there is and will remain a diverse range of production environments
globally. Indigenous breeds are adapted to the low-input to medium-input production
systems which account for about three quarters of production in the developing
world. However, encouraged by promises of high production, farmers are turning
to imported exotic breeds developed for high-input, comparatively benign agroeco-
systems. This displacement of locally adapted genetic resources often results in
reduced productivity and sustainability and higher risk because the exotic breeds
cannot tolerate the climatic, disease, and feed constraints and the frequent seasonal
swings in these components. Erosion of adapted genetics threatens not only food
security and the ability to maintain sustainable agriculture throughout all production
environments but also the cultural value of these indigenous resources.
The best livestock development strategy for sustainable food and agriculture
production and rural development is to improve or enhance breeds adapted to specific
production environments.
EVOLUTION OF THE GLOBAL STRATEGY
The Food and Agricultural Organization (FAO) mandate, which is regularly

reviewed by its 174 member governments, covers (1) collating, analyzing, and
reporting information; (2) providing technical assistance, with emphasis on the
developing world; and (3) providing a forum for intergovernmental discussion and
policy development. The member governments have resolved that FAO should lead,
coordinate, facilitate and report on the Strategy for the Management of Farm Animal
Genetic Resources (hereafter termed the Strategy). The term management incorpo-
rates the spectrum from surveying and characterizing, through to the development,
use, maintenance, and access to these resources. Member governments have accepted
a framework for the Strategy, and implementation involving donors and other stake-
holders has commenced. The implementation of the Strategy involving all stake-
holders is known as the Initiative for Domestic Animal Diversity.
Since the early 1980s, FAO has provided technical assistance to member coun-
tries in identifying and assessing their farm animal genetic resources, and in con-
servation and utilization activities. These activities, however, tended to be breed and
country specific and on a scale which bore little relation to the magnitude of the
problem. The organization did contribute to the development of practical method-
ologies for characterization and maintenance of animal genetic resources, and to a

growing awareness of the urgency and importance of addressing the problem of
biodiversity erosion on an international scale. While interventions ultimately must
be at the country level, farming systems and domestic animal breeds, and the threats
to their diversity and interest in accessing them, often cross national boundaries.
Further, an international approach to the management of this natural capital also
offers many opportunities for economies, for understanding and sharing the task,
and for the development and application of methodologies.
Major developments, such as the UN Conference on Environment and Develop-
ment (UNCED) in June 1992, the signing of Agenda 21, and ratification of the
Convention on Biological Diversity (CBD) in December 1993, reinforced the need
for the Strategy and served to shape the Strategy further. The Commission on
Sustainable Development mandated to review implementation of Agenda 21, and
the Conference of the Parties (COP) negotiating and implementing the articles of
the CBD will also influence the development of the Strategy.
In November 1996, parties to the Rome Declaration on World Food Security
recognized the need for sustainable food security and, to achieve this, for urgent
action on erosion of biological diversity, pledging to implement the World Food
Summit Plan of Action as their response. Also in November, the third meeting of
COP considered the Conservation and Sustainable Use of Agricultural Biological
Diversity as a major agenda item and decided in relation to AnGR that the Conference:
20. Appreciates the importance of the country-based Global Strategy for the Man-
agement of Farm Animal Genetic Resources under the Food and Agriculture Orga-
nization of the United Nations and strongly supports its further development,
(UNEP/CBD/COP/3/l.12)
PRINCIPLES OF AnGR AND THE RATIONALE

OF THE STRATEGY
Achieving “Food for All” will require sustainable intensification of agriculture

in many production environments. The genetic makeup of an animal is the key to
how it will respond to different aspects of the total production environment, partic-
ularly those aspects related to the uses demanded of the animal, to climate, feed and
water, exposure to disease, and to type of husbandry. This tenet must be accepted
to achieve and maintain sustainable farming systems, to realize production and
productivity increases, to manipulate product quality, and to minimize risk of pro-
duction losses over time. By definition, sustainability is specific to the production
environment. Further, there is and will remain a diverse range of production envi-
ronments globally.
Animals have been domesticated for thousands of years and have migrated over
time as human communities occupied new areas. Once settled in a new environment,
little further movement occurred but the local animal populations were changed
over time by the selection pressures exerted by the local environment, including the
imposition of human needs. The end result is that the gene pool of each farm animal
species has been redistributed such that about half the quantitative genetic differ-

ences are unique to any one breed while the other half is common to all breed
resources. It is these quantitative genetic complexes associated with adaptation and
performance that are central to food and agriculture production from animals.
Sustainable use of this between- and within-breed genetic diversity must be specific
to the production system.
While genetic diversity underpins current production, both in terms of the aver-
age level and consistency of production from season to season, diversity also provides
the basis for achieving the necessary improvements in production, productivity, and
product quality. Activities directed at such sustainable intensification of livestock
production should utilize genetic resources which are already adapted to the partic-
ular farming system production environments. This is because genetic adaptation
takes generations to develop, so that use of resource material which is already
adapted offers greater immediate and longer-term potential for achieving real and
sustainable gains in system performance. The challenge is to develop practical and
affordable means of realizing this goal in responding to the Commitments of the
World Food Summit Plan of Action (Rome Declaration).
The maintenance of genetic diversity also forms an essential hedge against future
threats to world food security, especially those posed by diseases, climate change,
and other modifications to the production environment. The rapid modernization of
agriculture, which has been so important in enabling the world to feed its expanding
population during this century, has promoted as the solution the development and
more or less universal adoption of high-intensity livestock production systems. These
systems have been based on a very few species and breeds that have been developed
to respond to intensive care, treatment, and feeding. Although less resilient to harsh
conditions, these developed breeds are displacing and contributing to the progressive
elimination of breeds which have been adapted over time to lower inputs and often
harsh environments. Although this widespread displacement is the greatest cause of
genetic erosion, there are several factors which place breeds at risk of loss and
threaten domestic animal diversity. These factors are listed in Table 1.
Table 1 List of Causes for Risk of Loss or Extinction of Breeds

Reason Description
Aid Lack of incentive to develop and use breeds, giving preference to those few
developed for use in high-input, high-output relatively benign environments
Product Undue emphasis placed on a specific product or trait leading to the rapid
dissemination of one breed of animal to exclusion and loss of others
Crossbreeding Indiscriminate crossbreeding which can quickly lead to the loss of original
breeds
Storage Failure of the cryopreservation equipment and inadequate supply of liquid
nitrogen to store samples of semen, ova, or embryos; or inadequate
maintenance of animal populations for breeds not currently in use
Technology Introduction of new machinery to replace animal draught and transport
resulting in permanent change of farming system
Biotechnology Poorly interpreted international genetic evaluation; artificial insemination and
embryo transfer leading to rapid replacement of indigenous breeds
Violence Wars and other forms of sociopolitical instability
Disaster Natural disasters such as floods, drought, or famine

Paradoxically, then, the very success of modern animal-breeding programs is the
principal factor redirecting effort from adapted and local breed use and development,
eroding the breadth of genetic material available for future breeding work.
A series of expert consultations supported by FAO and the United Nations
Environmental Program (UNEP) have identified and described the major issues
underlying the rationale for the Strategy,
1. Demand for animal products in most developing countries is increasing more

rapidly than for plant products; the main effect of this will be intensification of the
range of periurban and mixed farming production systems. This must be sustainable
intensification. It will also require that the large pastoral areas, to which local species
and breeds are well adapted, are kept in production under sustainable systems.
2. Increasing sophistication of developed country consumers is moving demand away
from uniform animal products to more varied products but with consistency of
each variant also being increasingly demanded. Animal welfare and human health
concerns are also being increasingly highlighted. Consumers are paying premium
prices for foods which are grown under lower-intensity systems and which meet
current lifestyle nutritional needs. These trends will continue to generate stronger
market incentives for the use of more-diverse genetic resources and consequently
for the maintenance of some additional farm animal biodiversity.
These observations reinforce the view that the most cost-effective way to maintain
animal breeds is the development of production and marketing policies that make
it financially attractive for farmers to maintain and improve local breeds. Sound
genetic resource utilization policy will be particularly important for the animal
species because of the high unit cost of individual animals and the often long
generation interval of species.
Preliminary survey results show that most countries possess a number of the
5000 or so remaining breeds of farm livestock, with the majority of these breeds
occurring only in developing countries. Most of these animal genetic resources are
owned by small farmers, emphasizing the importance of private good to the sound
management of these resources. The World Watch List for Domestic Animal Diver-
sity (FAO/UNEP, 1995) lists 3882 breeds for 25 domestic species in over 180
countries. Globally, 30% of breeds are classified as endangered and criticial based
on population size (Table 2). FAO defines “endangered” as populations having <1000
breeding females and <20 breeding males, and critical as populations having <100
breeding females and <5 breeding males. Of the breeds listed under these two
categories, 36% are managed either through a conservation program or maintained
by an institute. Presumably, the risk of loss of these breeds actively managed or
maintained is far lower than breed populations outside such management programs.
Of the total number of breeds with population data identified globally (2924), 19%
are categorized as endangered or critical and lack a breed conservation management
program. As such, there is a very high risk of loss of these animal genetic resources.
Where pricing systems and allocation of benefits are inadequate, public funding will
be required for effective management of these resources.
The development of competitive, sustainable production systems does not war-
rant the use of all existing breeds for a particular period of time. Consequently,

Table 2 Breeds of Domestic Animals at High Riska by Species
No. of Breeds
Categorized
as Critical or
No. Breeds Endangered
with (% of No. of Breeds
Total No. Population Which Are at High Risk of
Species Breeds on File Data ( of Total) Maintained) Loss (%)a
Mammalian
Ass 77 24 (31) 9 (0) 9 (38)
Buffalo 72 55 (76) 2 (0) 2 (4)
Cattleb 787 582 (74) 135 (41) 80 (14)
Goat 351 267 (76) 44 (16) 37 (14)
Horse 384 277 (72) 120 (20) 96 (35)
Pig 353 265 (75) 69 (25) 52 (20)
Sheep 920 656 (71) 119 (29) 85 (13)
Yak 6 6 (100) 0 ( 0 (0)
Dromedary 50 40 (80) 2 (0) 2 (5)
Bactrian camel 7 7 (100) 1 (0) 1 (14)
Llama 3 3 (100) 0 ( 0 (
Alpaca 4 4 (100) 0 ( 0 (
Guanaco 2 2 (100) 0 ( 0 (
Vicuna 3 3 (100) 0 ( 0 (
Total 3019 2191 (89) 501 (39) 364 (17)
Avian
Chicken 606 512 (85) 334 (61) 131 (26)
Turkey 31 29 (94) 11 (9) 10 (35)
Domestic duck 62 54 (87) 29 (38) 18 (33)
Muscovy duck 14 13 (93) 5 (40) 3 (23)
Domestic goose 59 51 (86) 28 (11) 25 (49)
Guinea fowl 22 17 (77) 4 (0) 4 (24)
Quail 24 23 (96) 16 (100) 0 (
Pigeon 19 16 (84) 2 (0) 2 (13)
Pheasant 8 7 (88) 0 ( 0 (
Partridge 11 4 (36) 0 ( 0 (
Ostrichc 7 7 (100) 3 (33) 2 (29)
Total 863 733 (85) 372 (48) 195 (27)
GLOBAL 3882 2924 (75) 873 (36) 559 (19)
a At risk determined based on breeds with population data having <1000 breeding females or
<20 breeding males and for which there is no conservation program in place.
b Includes mithan and banteng.
c Includes cassowary, emu, and nandu.
Source: Adapted from FAO/UNEP (1995).
utilization does not provide the universal solution for conservation of domestic
animal diversity. Nor is it essential to retain all breeds to ensure the ready availability
of the breadth of domestic animal diversity. Systematic breed-level characterization
of the genetic composition of species would make it possible to develop least-cost
strategies for maintaining a maximum of genetic breadth within each species from
a limited number of breeds.

Although semen and embryos of only a few of the endangered breeds are held
in cryopreservation storage, where the technology has been developed, it appears to
offer a viable alternative to live animal conservation when that is not feasible. The
nature of cryopreservation of animal cells renders such storage very high risk without
sample duplication.
The geographically dispersed nature of the resources and the need to involve a
range of stakeholders nationally and internationally strongly suggest that a cost-
effective Strategy must also incorporate efficient information-sharing systems. Mod-
ern data processing, storage, analysis, and communication technologies offer par-
ticularly important new potential for achieving this, and full advantage must be taken
of these opportunities in developing the Strategy.
Many policy issues concerning the good management of animal genetic resources
remain to be addressed, such as:
1. Ownership and sovereignty;

2. Access rights and sharing of benefits;
3. Facilitating use and development;
4. Roles of culture, gender, and indigenous knowledge;
5. Priorities for conservation of breed-level diversity;
6. Intellectual property rights;
7. Use of biotechnology in characterization, utilization, maintenance, and in research
and training;
8. Biosafety concerning the use of modified organisms and alien genetic resources;
and
9. Zoosanitary aspects of transport of resources for purposes of research, training,
and commercial use.
Because of the transboundary and legal implications which these present, such issues
must be dealt with by an intergovernmental mechanism; hence, the framework for
the Strategy must provide for an intergovernmental mechanism.
IMPERATIVES FOR ACTION
Based on this rationale for upgrading the management of farm animal genetic
resources, FAO has identified the following imperatives for action:
1. Identify and understand the genetic resources of each domesticated animal spe-
cies which collectively comprise the global gene pool for food and agriculture;
2. Develop and utilize correctly the associated diversity to increase production and
productivity, to achieve sustainable agricultural systems, and, where required, to
meet demands for specific product types;
3. Monitor, in particular, those resources which are currently represented by very
few animals or which are being displaced by breed replacement strategies;
4. Preserve the unique resources which are not currently in demand by farmers;
5. Train and involve people in all essential facets of management of these resources;
and
6. Communicate to the world at large the importance and value of the domestic
animal genetic resources and of the associated diversity, its current high exposure

Table 3 Working Definitions Proposed for Conservation
Animal Genetic Resources (AnGR) At the breed level, the genetically unique breed
populations formed throughout all domestication processes within each animal species used
for the production of food and agriculture, together with their immediate wild relatives (here
“breed” is accepted as a cultural rather than a technical term, i.e., to emphasize ownership,
and also includes strains and research lines).
Domestic Animal Diversity (DAD) The genetic variation or genetic diversity existing among
the species, breeds and individuals, for all animal species which have been domesticated and
their immediate wild relatives.
Conservation (of DAD) The sum total of all operations involved in the management of animal
genetic resources, such that these resources are best used and developed to meet immediate
and short-term requirements for food and agriculture, and to ensure the diversity they harbor
remains available to meet possible longer-term needs.
Conservation (in general) The management of human use of the biosphere so that it may
yield the greatest sustainable benefit to present generations while maintaining its potential to
meet the needs and aspirations of future generations. Thus, conservation is positive, embracing
preservation, maintenance, sustainable utilization, restoration, and enhancement of the natural
environment (IUCN-UNEP-WWF and FAO-UNESCO, 1980).
In Situ Conservation Primarily the active breeding of animal populations for food production
and agriculture, such that diversity is both best utilized in the short term and maintained for the
longer term. Operations pertaining to in situ conservation include performance-recording
schemes and development (breeding) programs. In situ conservation also includes ecosystem
management and use for the sustainable production of food and agriculture. For wild relatives
in situ conservation — generally called in situ preservation — is the maintenance of live
populations of animals in their adaptive environment or as close to it as practically possible.
Ex Situ Conservation In the context of conservation of domestic animal diversity, ex situ

conservation means storage. It involves the preservation as animals of a sample of a breed in
a situation removed from its normal production environment or habitat, and/or the collection
and cryopreservation of resources in the form of living semen, ova, embryos, or tissues, which
can be used to regenerate animals. While other methods of genetic manipulation, such as the
use of various recombinant DNA techniques, may represent useful means of studying or
improving breeds, they do not constitute ex situ conservation and may not serve conservation
objectives. At present, the technical capacity to regenerate whole organisms from isolated DNA
does not exist.
Source: Adapted from FAO (1995).
to loss, and the impossibility of its replacement. To facilitate effective communi-

cation a clear set of terminology is necessary. A minimum set of terms is given in
Table 3.
The Strategy must be country focused both to recognize properly the sovereignty
that nations have over their AnGR and because resources are lost or saved at the
country level. For success, and in understanding and developing genetic resources,
the Strategy must also focus interventions on identifiable production environments.
THE GLOBAL STRATEGY
To assist countries to respond to these imperatives for upgrading the management

of animal genetic resources, a framework for a global strategy has been supported
and is known as the Strategy for the Management of Farm Animal Genetic Resources.

The framework of the Strategy consists of four fundamental components. The
Strategy is designed to be comprehensive to emphasize the balanced approach
required to understand, utilize, and maintain AnGR better and more cost-effectively
over time. The four fundamental components of the Strategy are
1. A global, country-based structure comprising three elements:

a. Focal points and networks to assist countries design, implement, and maintain
comprehensive national strategies for the management of their AnGR. The need
has been clearly demonstrated to distribute the focal points to at least three
levels — country, regional and global — although for effective policy and
technical development subregional networks are also indicated for some
regions.
b. Stakeholder involvement, which FAO will lead and coordinate. This element
provides for a broad range of dimensions of involvement: geographical, AnGR
management element, production environment, species, time, and funding level.
c. A virtual structure to enable collection and use of information, coordination,
reporting, and to facilitate the spectrum of management processes. This virtual
structure is known as the Domestic Animal Diversity Information System
(DAD-IS) (Figure 1). It is being developed as an advanced communications
and information system primarily for country use. Its security protocols enable
countries to be responsible for validation of data and to determine what and
when data are released to the world, emphasizing the country-based concept.
DAD-IS also serves as the primary clearinghouse mechanism for this sector of
biodiversity as required under the CBD.
2. A technical program of activity aimed at supporting effective management action
at the country level, in harmony with the CBD, and comprising a set of six elements:
a. Characterization, encompassing demographic and environment, phenotypic,
and genetic indicators and assessment;
b. In situ utilization and conservation;
c. Ex situ conservation;
d. Communication and information system development, including the develop-
ment of DAD-IS and training;
e. Guidelines development and action planning; and
f. Collaboration, coordination, and policy instrument development (Figure 2).
3. Cadres of experts to guide development of the Strategy and maximize its cost-
effectiveness.
4. An intergovernmental mechanism for direct government involvement, policy devel-
opment, and support. This is provided by the Commission on Genetic Resources
for Food and Agriculture (CGRFA) (Figure 2).
To take full advantage of the Strategy, countries are being invited by FAO to
nominate an institution as a National Focal Point (NFP) and to identify a national
technical coordinator (NC). The NC serves as the point of contact for the involvement
of the country in the Strategy and will assist in organizing the essential in-country
networking, facilitating, and coordinating activity. To ensure the country level has
access to the necessary level of assistance and to best utilize the limited resources
of the Global Focus, the planned coordination structure provides for decentralization
to the regional level. Regional Focal Points (RFP) are to be implemented in each
major genetic storehouse region of the world.

Figure 1 Structure of Domestic Animal Diversity Information System (DAD-IS).
The technical program of activity is structured to facilitate implementation and

reporting and is being developed to align with the CBD (see Figure 2). Each technical
element provides for a set of connected activities, for example,
1. Characterization. Indicators and assessment of AnGR are essential to mount a

program of management. These are required at four levels:
a. Baseline surveys — national inventories of AnGR and of primary production
environments form the essential starting points for the development of Action
Plans, including the Early Warning System for AnGR.
b. Monitoring — review of the population status of particular AnGR at risk for
effective and efficient conservation and as the ongoing basic need for Early
Warning.
c. Comparative genetic evaluation — increased knowledge of the unique qualities
of breeds under each production environment, as the basis for making best use
of these resources in the short and longer term.
d. Genetic distancing — a comparative molecular description strategy is being
developed to establish which breeds harbor significant unique genetic diversity,
allowing better-targeted conservation actions and facilitating baseline surveying.
2. In situ utilization and conservation. These activities primarily involve the active
breeding of animal populations such that genetic diversity is best utilized in the
short term and maintained for the longer term. In situ operations include establishing

Figure 2 Primary intergovernmental and technical work elements of the Strategy and their
relationship with the CBD, CSD, and Agenda 21.
breeding goals for sustainable production systems and performance recording and
genetic development and dissemination of improved germplasm. In situ conservation
covers the maintenance of live populations of animals in their natural environment
for possible future use. Incentive systems will generally need to be developed to
enable farmers to develop and reliably maintain such conservation activities.
3. Ex situ conservation. This activity includes cryogenic preservation and the main-
tenance of live animals in farm parks or zoos, beyond their development environ-
ment. The Strategy focuses on the use of live animals backed up by
cryopreservation where technology exists, combining within-country genome
banks with likely regionally distributed global genome repositories of last resort.
For the latter in particular, policy development is required to assist countries to
secure diversity associated with resources currently at high risk.
4. Guidelines and Action Plans. Cost-effective management of AnGR is complex
technically and operationally. The Strategy provides assistance to countries through
the development and provision of comprehensive guidelines for use as decision
aids in the planning and implementation of national action strategies. These guide-
lines will form much of the detailing at the center of the Strategy; consequently,
their rapid development and field testing will best assist countries. The Strategy
will then be further developed by integrating all national action strategies.
FROM STRATEGY TO A GLOBAL INITIATIVE
The components of the Strategy are interdependent, and require concurrent

implementation to maximize cost-effectiveness, generate momentum, and guarantee

Figure 3 The Global Initiative for Domestic Animal Diversity involves all stakeholders.
success over time. Still, once the guidelines and basic infrastructure are in place,
involvement must be developed over time, as human and financial resources become
available.
The collective collaborative effort being developed to implement the Strategy
is termed the Initiative for Domestic Animal Diversity (iDAD). The Initiative com-
prises all stakeholders: the countries themselves — their farmers, governments, and
focal points; financial institutions; international agencies; and nongovernmental
organizations, including the commercial sector. Figure 3 summarizes the Initiative.
An informal mechanism for stakeholder involvement is being developed utilizing
both real and virtual conferencing to achieve the range and continuity of involvement
required. Building upon the Strategy, the key objectives of the Initiative are sum-
marized in Table 4.
Table 4 Objectives of the Initiative for Domestic Animal Diversity

• Support and develop a global, country-based structure to assist countries;
• Help countries design and implement action plans and management strategies;
• Promote a better understanding of farm AnGR and the threats to them;
• Facilitate the use and improvement of locally adapted genetic resources;
• Establish the comparative uniqueness of resources in order to maximize the cost-effectiveness
of their management;
• Enable the maintenance of unique AnGR by assisting development of national and global
genome banks;
• Ensure the Strategy conforms to the CBD;
• Stimulate and coordinate the involvement of the range of stakeholders;
• Promote and enable intergovernmental discussion and policy development;
• Promote development and use of better technologies for cost-effective strategies;
• Upgrade training and facilitate capacity building;
• Be aware of the status of the world domestic animal diversity; and
• Facilitate access to irreplaceable animal genetic resources.

Table 5 Species of Livestock Receiving
Initial Focus in the Global Strategy
Ass Domestic goose
Buffalo Horse
Camel Lamoids
Cattle Pig
Chicken Rabbit
Duck Sheep
Goat Turkey
The Strategy will be implemented over 15 years, progressing through the phases
of implementation from awareness and development of a global structure for coor-
dination, to assisting countries implement national action strategies. The FAO gov-
erning body has requested the focus initially be on the 14 or so most important
livestock species which account for over 90% of agriculture and food production
(Table 5).
As the UN technical secretariat for agriculture, FAO is uniquely placed to
facilitate, coordinate, and report on AnGR. FAO is providing some essential core
resources for the operation of the Global Focus for the iDAD. However, it is neither
possible nor appropriate for FAO to provide all the human, financial, and other
resources required for effective implementation of the Strategy.
The Strategy aims to increase awareness and understanding of AnGR, markedly
improve their use and sustainable development, upgrade the maintenance of unique
resources, and enable improved access and benefit sharing. It will promote interna-
tional collaboration and markedly increase the cost-effectiveness of AnGR-related
initiatives worldwide, avoiding duplication of effort. The Strategy provides for a
range of bilateral and multilateral assistance and for public and private sector support.
It is important to note that such support need not necessarily be through FAO.
Importantly, the Strategy provides for countries to discuss and develop directly the
broad range of policy issues concerned with the critical upgrading now required for
the management of AnGR of interest to food and agriculture, with their improved
characterization, use, development, maintenance, and access.
BENEFITS OF THE GLOBAL STRATEGY
Based on compelling evidence and experiences, specific benefits associated with

the Strategy include the following:
1. Framework for country assistance and action planning:

a. A structural framework to assist countries manage their AnGR;
b. A mechanism for country decision making and international policy develop-
ment;
c. An integrated and comprehensive technical framework to provide direction for
national effort, promote expert comment and advice prior to expensive imple-
mentation of action plans, and give sound guidance for the maintenance of
action strategies; and

d. Increased country and regional institutional, managerial, scientific, and techno-
logical capacity to improve productivity of AnGR and maintain domestic animal
diversity.
2. Development, sustainable use, and maintenance of Animal Genetic Resources:
a. Coordinated surveying, monitoring, and description of AnGR;
b. Establishment of the comparative genetic uniqueness of AnGR to minimize
management costs over time;
c. Improved development, use, and maintenance of AnGR according to a strategic
combination of in situ and ex situ conservation and utilization;
d. Better matching AnGR to feed resources and other inputs, and to required
outputs, to increase productivity, reduce environmental degradation, and main-
tain sustainable production systems; and
e. Conservation of domestic animal diversity of national and global importance,
retaining for future generations the necessary broad flexibility required to
respond quickly and efficiently to unpredictable change in production environ-
ments.
3. International collaboration and policy development:
a. A mechanism to assure continuity and coordination of AnGR-related activities
over time;
b. A strong project identification mechanism to propose costed projects and activ-
ities for financing and to minimize duplication;
c. Detailed country national action strategies to enable donors to recognize invest-
ment priorities; and
d. Through the CGRFA, a mechanism for intergovernmental deliberation and
policy development.
4. A communication and information tool, DAD-IS
a. Has been developed as the key communications and information tool of the
Strategy for country use, to link and involve all other stakeholders with the
Strategy;
b. Incorporates security features to enable countries to use the system as their
own, with complete control over national data and other information;
c. Provides databases for cataloging domestic animal breeds, their characteristics
and risk status, production environments, and the range of activities underway
for each breed, thereby acting as a comprehensive information system for AnGR
use, development, access, and early warning;
d. Enables countries and international collaborators to set conservation priorities;
e. Supports utilization and conservation by communicating the values and roles
of AnGR;
f. Enables global networking by putting countries and their farmers in contact
with other farmers; research, training, and education institutions; and scientists
from around the world;
g. Provides guidelines for development of national action strategies and supports
action planning for the sustainable development, use, and maintenance of
AnGR;
h. Supports cost-effective training and research; and
i. Provides customized project management software to support AnGR activity
planning, reporting, and monitoring.
A comprehensive communications element for the Global Strategy will support
all stakeholders and provide a least-cost system for awareness generation at all
levels.

5. CBD objectives:
The components and elements of the Strategy are being developed to align with
the objectives of the CBD: conservation, sustainable use, and sharing of the
benefits of biological diversity.
6. World Food Summit Plan of Action:
For the farm animal biodiversity important to developing “Food for All” on a
sustainable basis, the Strategy provides an integrated framework for action,
particularly with respect to Commitments Three and Seven of the Plan, which
refer to signatory governments pursuing participatory and sustainable develop-
ment policies in high and low potential areas, and implementing, monitoring,
and following up the World Food Summit Plan of Action at all levels.
The preceding benefits represent a powerful incentive for implementing a strong

Strategy for the Management of Farm AnGR.
STRENGTHS AND INNOVATIVE FEATURES OF

THE GLOBAL STRATEGY
The benefits are secured through a number of strengths and innovative features
of the Strategy, which
1. Facilitates assessment of AnGR and incorporates development of a comprehensive

early warning system, providing the basis for establishment of conservation prior-
ities and action plans;
2. Provides the necessary framework for a participatory approach to building country
consensus around complex issues, for resolving conflicting situations, and devel-
oping effective policy;
3. Incorporates a specific mechanism to facilitate country, subregional, regional, and
other international cooperation in AnGR management;
4. Offers a unique mechanism to identify, link, and strengthen the network of regional
and national institutions, based in the main on existing structures and their involve-
ment in the development of the Strategy;
5. Provides a framework for assistance that promotes clear identification of priorities
and links country, regional, and global priorities and concerns to maximize the
impact of funded activities;
6. Makes available a unique moderated, comprehensive information and communi-
cations tool — DAD-IS — which can be used by countries as their own secure
system while also enabling all stakeholders to be informed and involved; this new
system also provides the primary clearinghouse mechanism for the farm animal
sector of agrobiodiversity;
7. Provides the capability for ongoing project and activity documentation and moni-
toring by countries and donors, and for least-cost formulation of action plans at the
species, country, regional, and global levels; through the DAD-IS Action Planner;
8. Promotes, for this sector of biological diversity, the development of a management
approach which is in harmony with the CBD and in pursuance of Agenda 21;
9. Comprehensively provides for the involvement of the spectrum of donors and other
stakeholders;
10. Allows for coordination of effort while respecting stakeholders’ need for control
of their own activity; and

11. Incorporates guidance from the necessary disciplinary array of technical specialists
convened regularly to develop the Strategy further and advise on its implementation.
While the Strategy has been supported and early implementation has commenced,1
success depends on its ability to mobilize the commitment, energies, and resources
of the many different groups which have a stake in conserving and utilizing agro-
biodiversity — farmers and pastoralists throughout the world, those involved in
training and research into improved farming systems and technologies, environmen-
talists, agroindustries, and ultimately the consumers. While countries will be for-
mally responsible for developing and implementing national policy, supported by
the international collaborative effort, implementation of the Strategy must involve
these additional stakeholders and be responsive to their varied concerns, creating
mechanisms for interaction.
REFERENCES
FAO/UNEP, 1995. World Watch List for Domestic Animal Diversity, 2nd ed., FAO, Rome.
Hammond, K. and Leitch, H., 1996. The FAO Global Programme for the Management of
Farm Animal Genetic Resources, in Beltsville Symposia in Agricultural Research XX,
Biotechnology’s Role in the Genetic Improvement of Farm Animals, R. H. Miller, V. G.
Pursel, and H. D. Norman, Eds., American Society of Animal Science, Savoy, IL, 24.
IUCN/UNEP/WWF and FAO/UNESCO. 1980 World Conservation Strategy, Living
Resources Conservation for Sustainable Development, IUCN, Switzerland.
1A listing of activities underway and progress to date implementing the Strategy is available on the
Internet at http://www.fao.org/dad-is.

CHAPTER 16
Agroecosystem Quality: Policy and

Management Challenges for New
Technologies and Diversity
Joel I. Cohen
CONTENTS
Introduction
Confronting the Diagnostic Challenge: Technical vs. Adaptive Problems
Introducing Agroecosystems and Indicators of Quality
Defining Agroecosystems
Factors Affecting Quality Indicators
Quality Indicators — Linking Biodiversity with New Technologies
Conserving, Maintaining, and Using Biodiversity
Minimizing Chemical Inputs
International Collaboration in Biotechnology Research
Findings
Anticipating Adaptive Challenges for Developing Countries
Seminar Findings
Examples from IBS Seminars: The Technical and Adaptive Challenges
The Case of Durable Resistance to Rice Blast Fungus
The Case of Bacillus thuringiensis and Transgenic Crops
Quality Indicators and New Technologies — Synthesis of Above
Discussion
Agroecosystem Quality and Challenges Ahead — Adaptive Problems
Revisited
References

INTRODUCTION
Providing a meaningful contribution to the topic of agroecosystems, new tech-

nology, and diversity poses many challenges. First, it is difficult to obtain agreed-
on definitions or standards for “agroecosystem quality.” The second difficulty occurs
when considering how new technologies affect agroecosystem quality, including
issues related to biodiversity. These difficulties, and the management and policy
issues which they raise, are illustrated by examples of technical and adaptive chal-
lenges facing agricultural policy makers, managers, and end users concerned with
maintaining levels of biodiversity or enhancing agroecosystem quality.
The objectives of this chapter are to first consider the differences between these
technical and adaptive problems, the nature of the situations they each address, and
the learning required when facing an adaptive challenge. Second, agroecosystem
complexities and the difficulties in determining quality indicators are presented.
Applications of biotechnology are presented as derived from international collabo-
rative research using examples compiled by the Intermediary Biotechnology Service
(IBS), executed by the International Service for National Agricultural Research
(ISNAR). Some of these examples, as used in IBS policy seminars, highlight
emerging policy and management needs which were identified and discussed. It is
hoped that this chapter clarifies adaptive challenges regarding agroecosystem diver-
sity and quality, and prepares stakeholders for the challenges and opportunities of
new technologies.
CONFRONTING THE DIAGNOSTIC CHALLENGE:

TECHNICAL VS. ADAPTIVE PROBLEMS
When confronting “technical problems,” difficulties are faced which can be

clearly defined and understood, and for which solutions are readily available. They
have become problems of a technical nature by virtue of lessons learned through
experiences confronted over time. The benefits derived from these accumulated
experiences let us know both what to do, through the use of knowledge (organiza-
tional procedures for guiding our actions), and who should do it, by identifying
whoever is authorized to perform such work (Heifetz, 1996).
When facing an “adaptive problem,” however, ready organizational responses
are absent, the problem is difficult to define, and expertise and/or established pro-
cedures are lacking. Technical responses to the problem are at best only part of the
solution. When facing such difficulties, time is required for learning, as this is a
central task of the adaptive process. Learning occurs before solutions and imple-
mentation modalities become apparent. Those holding competing values with regard
to the problem are identified, questions are posed to define the issues, and stake-
holders are given time to adjust values to accommodate the nature of the problem.
The learning phase of adaptive work diminishes the gap between the original stake-
holder values, the realities they now face, and the adjustments that may be necessary
to adapt their values to the new realities (Heifetz, 1996).

Differences between technical and adaptive problems are used to diagnose issues
presented in this chapter as related to agricultural productivity (see Table 1). Agri-
cultural problems of a technical nature are often remedied by choosing among
appropriate technologies, whether they are from conventional or nonconventional
sources. One chooses between or combines various cultural, crop, or livestock
options to address problems, needs, or deficiencies in productivity of agricultural
ecosystems. However, when technologies are considered beyond their technical
dimensions, in the broader sense of affecting agroecosystem quality, then adaptive
problems may be encountered for the following reasons. First, no universal definition
of quality exists, especially for the variable nature of agricultural ecosystems in the
tropical climates of developing countries. Second, stakeholder opinions may vary
as to utility vs. risk of new inputs or technologies. Third, values (whether cultural,
economic, or health) create perceptions which must be addressed in relation to the
realities of the proposed inputs and the changes they may cause. It is in this context
that new technologies can raise adaptive challenges to farmers, system managers,
and policy makers.
Consequently, questions regarding agroecosystem quality are “adaptive chal-
lenges.” In this paper, two indicators of agroecosystem quality are proposed, one
based on biodiversity and the second on the use of chemical inputs. These indicators
can be affected by the introduction of new technologies, using biotechnology
products as examples. Biological differences among agroecosystems and stake-
holder values and perceptions will be critical to defining specific quality indicators.
Policy and management challenges posed by new technologies and considerations
of biodiversity and use of chemical inputs are then analyzed in relation to agroec-
osystem quality.
INTRODUCING AGROECOSYSTEMS AND

INDICATORS OF QUALITY
Defining Agroecosystems
Agroecosystems include highly managed, productivity-oriented systems which

vary widely in their dependence on chemical, energy, and management inputs, and
are one conservation tactic identified to protect extant diversity (Soule, 1993). Defin-
ing “quality indicators” associated with agroecosystems relies on concepts not inher-
ent in the system itself, just as do efforts to define sustainability. Rather, concepts
such as sustainability or “quality” imply values derived from a human or cultural
perspective for a particular management system (J. Tait, personal communication).
These perspectives help determine whether a particular agricultural input enhances
agroecosystem quality or not.
Four major components of agricultural systems have been proposed by Antle
(1994) in studies on pollution and agriculture. His work highlighted relations among
(1) agricultural production, (2) the broader agroecosystem, (3) human health con-
siderations, and (4) valuation and social welfare, with each possessing characteristics

Table 1 Summarizing the Technical and Adaptive Problems, Solutions, and Questions Related to Agroecosystem Quality, Biodiversity, and New
Technologies
I.A Technical problems • Clear problem definition
characterized by: • Clear problem solution
• Able to identify relevant authority/developer for solution
I.B Technical problems and Problem 1: Is durable resistance available for rice blast in farmer’s
solutions posed: fields?
Technical Solution: Improved varieties, with new sources of genetic resistance
Problem 2: Is insect resistance using B.t. available in tropical maize?
Technical Solution: Improved varieties, with new sources of genetic resistance
II.A Adaptive problems • Organizational responses are absent,
characterized by: • The problem is difficult to define,
• Expertise and/or established procedures are lacking
• Technical responses are at best only part of the solution
• Time required for learning
II.B Adaptive problem posed in Does the introduction and use of described products Two indicators of quality selected in this paper:
this paper: require changes in stakeholder values, perceptions, • Biodiversity, conservation and use
or attitudes with regard to agroecosystem quality? • Minimize use of chemical inputs
III Answers depend on ability to In the view of the stakeholders:
address questions, such as: • Have new sources of resistance affected the composition of extant biodiversity, including possibility for horizontal
gene transfer?
• Have the new varieties diminished the need for chemical insecticides or fungicides?
• Have new varieties included management packages for gene deployment, and extending or guarding the length
of time available for resistance?
• Are clear understandings available for current chemical input levels?
• Are measures of productivity or other economic gains available?
• Was the technical problem solved?

valued by society. By using the divisions presented by Antle, the introduction of
novel sources of genetic diversity would occur in the agricultural production. Cou-
pling the introduction of biotechnology with the management of biodiversity and
agroecosystem quality would influence a range of perspectives regarding overall
quality of the agroecosystem component (2) and, often, values of human health and
welfare (3 and 4).
Factors Affecting Quality Indicators
Determining practices to enhance the sustainability of a given agricultural sys-

tem, as presented by Tait (personal communication), and the components used by
Antle (1994) in his pollution study are also useful for this discussion. Here, these
two concepts (dependence on human values and four components depicting intro-
ductions to agricultural systems) are used in the context of managing agroecosystems
in developing countries. They provide a foundation for understanding the interrela-
tions between quality indicators, inputs derived from biotechnology, and agroeco-
system biodiversity. Examples of inputs are given, using cultivars as technical solu-
tions to specific environmental and productivity problems, but which can also be
valued in the context of the ecosystem.
QUALITY INDICATORS — LINKING BIODIVERSITY

WITH NEW TECHNOLOGIES
Relevant agroecosystem quality indicators, which could be applied to products

derived from new technologies, now need to be selected. Examples of products, like
virus resistance and applications of B.t. (see section on Examples from IBS Seminars,
later), illustrate both technical and adaptive challenges when considered in relation
to agroecosystem quality. With such examples in mind, two indicators were selected
which would relate them to agroecosystems: (1) biodiversity and (2) diminishing
use of chemical inputs.
Conserving, Maintaining, and Using Biodiversity
Many traditional agroecosystems are undergoing some process of modernization

(Altieri and Merrick, 1988). This process of modernization and its relation to the
use of high-yielding varieties can threaten indigenous diversity or other repositories
of crop germplasm. Pressures to modernize can have a drastic effect on the conser-
vation of diversity, and indicators of quality will depend on our knowledge of natural
populations in each ecosystem. In many agroecosystems, premiums are placed on
maintaining and conserving sources of biodiversity. Different and often competing
values exist for what constitutes an ecologically correct mix or use of diversity within
a given agroecosystem. Whether this diversity can be increased or decreased reflects
values attributed to ecosystem quality. Placing premiums on maintaining diversity
recognizes the importance of multiple-crop agroecosystems which make use of
indigenous as well as introduced sources of diversity (Gliessman, 1993). Complex

crop mixtures, rotations, and practices developed by local farmers can protect the
environment under tropical conditions and provide an array of products for harvest.
Several case study examples illustrate the importance of using and conserving
extant biodiversity within managed agricultural and forest ecosystems (Potter et
al., 1993). An important, if not essential, element of these systems is the involvement
of native peoples in these managed areas, and their application of the knowledge
gained over time for the care and management of such areas (Padoch and Peters,
1993). In addition, it has been argued that maintaining traditional agroecosystems
is an important strategy for preserving in situ repositories of crop germplasm (Altieri
and Merrick, 1988). For example, Latin American farming systems studied dem-
onstrate a high degree of plant diversity (Altieri and Montecinos, 1993). The authors
also recognize the importance of small farmer holdings in these ecologically diverse
systems.
Minimizing Chemical Inputs
Biotechnology and sustainable agricultural systems are often portrayed as antag-

onistic ends of a continuum. However, this portrayal lacks evidence, especially given
that the use of biotechnology-derived agricultural products within either production
systems or agroecosystems is still largely an unknown factor. In fact, there are many
applications of biotechnology which seek to minimize the use of chemical inputs
as pest, weed, or disease control strategies in developing country agriculture. The
relation between these applications and broader concerns of sustainability have been
recognized (Hauptli et al., 1990). In this regard, technical solutions to pressing pest
or weed management problems are becoming available from biotechnology. For this
reason, minimizing chemical inputs to agroecosystems was selected as the second
potential quality factor to be presented.
Both of these indicators will rely on mobilizing, understanding, and taking into
account stakeholder values and perceptions. Management of agricultural systems
will be complicated by the fact that indicators of quality are difficult to measure,
highly location specific, and reflect “value judgments.” Such indicators will by
necessity incorporate values held or determined by the stakeholders of each system,
and will reflect values that are not part of the biological system being considered
(J. Tait, personal communication). Solutions to stakeholder problems, such as the
need to combat pests or minimize chemical applications, can take the form of
technical solutions by using new inputs. However, adaptive problems may also occur
after interventions are identified and new technical solutions are employed. Here,
stakeholder opinions may differ with the claims made by or for technical solutions,
such as can occur with new products from agricultural biotechnology, or when levels
of extant diversity are threatened.
It is necessary to identify the real stakeholders, to learn their expectations
regarding the issue, and to gain an understanding of their opinions regarding these
options to the problem at hand. Mobilizing stakeholder response is a key facet of
adaptive problems, and a major task for those managing such situations (Heifetz,
1996). Constituents of specific agroecosystems will help determine quality indicators
and work with those advocating new inputs, or cultural options which may affect

levels of diversity. Introducing new sources of diversity raises further complications
in agreeing whether such additions reflect an improvement in overall quality. These
complications are expected, based on the increases in stakeholder involvement
regarding the question of genetically engineered crops and introductions to areas
rich in extant or indigenous biodiversity.
INTERNATIONAL COLLABORATION IN BIOTECHNOLOGY RESEARCH
With the two indicators of agroecosystem quality determined, attention is now

placed on examples of new technologies. Examples have been selected that take into
account the emerging needs of developing countries regarding biotechnology and
their ability to collaborate with international research programs. These examples are
taken from information collected from IBS policy seminars and its Registry of
Expertise. IBS began to collect, analyze, and discuss with client countries its infor-
mation on international collaboration in biotechnology by organizing a meeting held
at ISNAR in 1993 (Cohen and Komen, 1994).
Information was collected through survey forms from some 40 international
biotechnology programs. Taken together, this material clearly demonstrated that
international collaboration in agricultural biotechnology offers developing countries
access to a range of specific technologies, and unique opportunities for developing
improved crop plants, livestock, vaccines, and diagnostic probes. An aggregate
analysis of this information was made, as described below, for which specific
conclusions are most relevant for a discussion on new technologies and agroecosys-
tem quality.
Findings
Among the international programs studied by IBS, most research is undertaken

on essential commodities, or foods on which significant numbers of people depend,
often with regional significance (Brenner and Komen, 1994; Cohen and Komen,
1994; IBS, 1994). Analysis of the 22 international crop biotechnology research
programs indicates that they address five broad research objectives, containing 126
separate activities. These primary objectives, crops, and research activities are shown
in Table 2. As such, they represent solutions to many technical situations facing
farmers and growers in developing countries.
With regard to crop transformation, research supported by the international
programs concentrates primarily on resistance to viruses and insects, and improving
quality factors (IBS, 1994). In Table 3, general categories and specific examples of
transformation are shown for agriculture in industrialized countries, using examples
from Day (1993). The third column summarizes research being conducted specifi-
cally for developing country agriculture with illustrations of specific applications.
These data indicate a strong commitment to improving crop plants through
biotechnology by addressing agricultural needs and objectives for developing coun-
tries. Approximately 50% of the expenditures in these international biotechnology
programs are devoted to research needed to develop these modified crops (Cohen,

Table 2 Number of Research Activities Undertaken by International Biotechnology Projects
as Shown for Five General Research Objectives and for Crops of Major Importance
to Developing Countries
Objectives
Disease Insect Virus Quality
Crops Resistance Resistance Resistance Traits Micropropagation All
Cereals 9 13 8 12 42
Rice 5 4 6 6 21
Maize 1 6 2 3 12
Sorghum 1 3 2 6
Other 2 1 3
Root Crops 4 5 7 2 1 19
Potato 1 3 2 6
Cassava 1 3 2 6
Yam 2 1 1 4
Sweet potato 2 1 3
Legumes 4 6 4 6 20
Bean 1 2 1 2 6
Groundnut 1 1 3 1 6
Chickpea 1 1 2 4
Other 1 2 1 4
Horticulture 2 3 1 6
Perennial 2 1 2 2 15 22
Banana/plantain 2 1 5 8
Industrial crops 1 4 5
Coffee 1 4 5
Sugarcane 1 1 1 3
Cocoa 1 1
Forestry Species 2 5 7
Miscellaneous 3 3 2 2 10
All 24 28 24 26 24 126
Note: Figures are based on information gathered from 22 international research programs that
include activities in crop research. For this table, we used those research activities with a
specific applied objective, excluding research activities aimed toward general technology
development.
From IBS BioServe Database, 1997.
1994). This percentage of available resources increases their ability to solve technical
problems, as defined in this chapter, and as shown in the examples below. However,
this also means that a much smaller amount of resources is available to address
questions of a more adaptive nature arising as their products move from research
into agricultural production, and then enter the broader agroecosystem, confronting
human health or valuation considerations (Antle, 1994).
Anticipating Adaptive Challenges for Developing Countries
Over the past 4 years, IBS has organized a series of Agricultural Biotechnology
Policy Seminars, held regionally for collaborating countries. In these seminars,
attention is given to examples of biotechnology providing solutions to technical
problems faced by farmers in developing countries. These same examples are

Table 3 Cloned Genes of Interest for Crop Plant Improvement and Related Applications
of the International Biotechnology Programs
International
Biotechnology Program
General Categorya Specific Examplesa Applicationsb
Disease resistance: viruses Virus coat protein subunits African cassava mosaic virus,
(TMV, cucumber mosaic, common cassava mosaic
potato virus X) virus
Potato leaf roll virus Bean gemini viruses
Potato virus S Rice stripe virus, yellow
Soilborne wheat mosaic virus mottle virus, tungo virus,
Plum pox virus ragged stunt
Tomato spotted wilt virus Potato virus X and Y
Viral replicase gene (PVX) Tomato yellow leaf curl virus
Sweet potato feathery mottle
virus
Groundnut stripe virus,
Rosette virus, and clump
virus
Fungal diseases Chitinase gene, H1 gene for Potato late blight
resistance to H. carbonum Rice blast
from maize, systemin gene
— a peptide signal molecule
which controls wound
response in plants,
infectious viral CDNA
Insect resistance B.t. genes, cowpea trypsin B.t. toxin genes applied to
inhibitor, wheat agglutinin borers in maize, rice,
gene for resistance to sugarcane, potato, coffee
European corn borer Potato glandular trichomes
Sweet potato weevil
Pigonpea: Helicoverpa and
podfly
Storage protein genes Wheat low-molecular-weight No applications reported
glutenin gene, maize
storage protein
Carbohydrate products Polyhydroxybutyrate as an No applications reported
alternative to starch for the
production of biodegradable
plastics
Ripening Antisense polygalaturonase No applications reported
in tomato, regulation of ACC
synthase gene
Breeding systems Self-incompatibility genes Male sterility in rice
from Brassica, anther
specific genes used for male
sterility with a ribonuclease
gene
Flower color Petunia, Antirrhinum No applications reported
Herbicide resistance Glyphosate, bialaphos, and No applications reported
imidazolinone resistance
a General categories and specific examples from Day, 1993.
b Examples from IBS (1994) BioServe database of international agricultural biotechnology
programs.

explored with regard to the adaptive challenges posed when new technologies enter
agricultural systems. As in many complex social situations, agricultural managers
and policy makers can face substantially more complex adaptive challenges from
situations originally perceived as technical in nature. Often, the problem itself is
unclear because of divergent opinions regarding the nature of the problem and its
possible solutions (Heifetz, 1996). One stakeholder’s technical solution is another
stakeholder’s adaptive challenge. In these cases, there is also often disagreement
among scientific experts, particularly at early stages of problem definition, hence
the time needed for learning.
In the seminars, technical examples are explored from the perspective of multi-
disciplinary and diverse national delegations. In facilitating these delegations, IBS
ensures involvement of individuals with responsibility for, or vested interest in, the
design, implementation, and use of agricultural biotechnology. This range of stake-
holder interests enriches the debates which occur within each delegation as the
delegates identify needs for services to help with the learning phase of adaptive
work, often taking the form of policy dialogues, management recommendations, or
responses needed for various international agreements. As such, IBS builds on
scientific data and available understanding to expand discussions to address the
broader needs of stakeholders, including policy makers, managers, and researchers,
and farmers, end users or non-governmental organizations (Komen et al., 1996).
Seminar Findings
Participant action planning methodology, carried out by the 17 attending coun-

tries, identified needs and/or constraints. In total, 227 needs were identified from
the delegations. These needs were systematically analyzed, identifying nine general
policy issues, their relative degree of emphasis, and whether or not there was a
convergence of these needs (Table 4). In addition, seven implementation issues and
three issues related to priority setting have been summarized. Most relevant to a
discussion on new technologies and agroecosystem diversity are the needs identified
for biosafety, socioeconomics, and priority setting. Here, the specific needs related
very clearly to the adaptive policy challenges facing developing countries, particu-
larly those located in centers of diversity. These issues will be presented later, in the
section on Quality Indicators and New Technologies.
EXAMPLES FROM IBS SEMINARS:

THE TECHNICAL AND ADAPTIVE CHALLENGES
In the most recent policy seminar for selected countries of Latin America, three
case studies were presented on issues related to biotechnology, productivity, and the
environment. These case examples are most relevant to the discussion above. They
illustrate solutions to agricultural problems having, to a greater or lesser extent, an
adaptive and technical component (Roca et al., 1998; Serratos, 1998; Whalon and
Norris, 1998).

Table 4 Number of Policy Needs Identified by Members of 17 National Delegations
Attending Policy Seminars for Africa, Asia, and Latin America
No. of
No. of Countries No. of Needs Convergent
General Policy Issues Responding Identified Needs
1 Biosafety 14 19 4
2 Socioeconomic assessment 12 19 3
3 Integration 9 11 2
4 Policy development/coordination 9 9 2
5 End user/beneficiary linkages 9 10 2
6 Technology transfer system 8 8 2
7 Intellectual Property Rights (IPR) 7 8 3
8 Biodiversity 6 7 3
9 Public awareness 5 5 1
The first example uses the introduction of improved rice varieties with the
potential to curtail use of toxic and expensive fungicides. This case is primarily
technical, as the products and techniques used have not posed adaptive challenges.
In this case, the new varieties are not products of transgenic technologies. Rather,
biotechnology tools have been used after varietal development to understand sources
of resistance and to type resistance against lineages of the pathogen. For the second
case, the introduction of maize containing novel sources of resistance to insect pests
is considered. In the case of maize, insect resistance is derived from transgenic
technologies allowing for the insertion of genes encoding a pesticide from bacteria.
In the third case, broader implications of managing and deploying transgenic crops
using Bacillus thuringiensis (B.t.) technologies are considered. As can be seen from
the maize and the B.t. examples, complex situations can be anticipated when intro-
ducing new inputs into traditional agroecosystems.
The Case of Durable Resistance to Rice Blast Fungus
Blast is the most widespread and damaging disease of rice. When control is needed,
and is not present in the form of cultivar resistance, then fungicide treatments are
applied which may not be effective, economically sound, or desirable from an envi-
ronmental perspective. Conventional resistance has been made available genetically,
but it has traditionally been weakened or lost after 3 years. However, durable resistance
has been achieved in rice cultivars using conventional breeding, resulting in Oryzica
Llanos 5, developed as a resistant variety by Centro International de Agricultura
Tropical (CIAT), the National Federation of Rice Growers, and the National Research
Institute of Colombia (F. Correa-Victoria, personal communication).
The variety was introduced to tropical agroecosystems in Colombia and repre-
sented a technical solution to the problem of blast, as well as the potential to improve
system quality by reducing the unwise or ineffective use of fungicides. The cultivar
was adopted across Colombia in the season following its release, and has been
planted in at least 50,000 ha/year until 1996. Since then, newer high-yielding cul-
tivars were released and widely adopted by farmers (F. Correa-Victoria, personal
communication).

More recently, techniques derived from biotechnology have been coupled to
these applied breeding strategies (Tohme et al., 1992; Roca et al., 1998). These
molecular tools are helping to understand the mechanisms controlling durable resis-
tance in Oryzica Llanos 5 by typing resistance genes to different genetic families
of blast, identifying molecular markers associated with resistance genes in other
highly resistant cultivars, and guiding rice breeders in selection of potential parents
leading to lines with durable blast resistance. Genes are being identified that express
resistance to six pathotype lineages of the blast pathogen. This analysis depended
on the use of DNA probes containing cloned fragments of the blast fungus genome
which could then be used to construct DNA fingerprints of the fungus. Molecular
markers were then used by breeders to confirm the manipulation and selection of
various sources of resistance to these six lineages of the blast fungus. This resistance
will bring a reduction in the use of fungicides by farmers as in the case of the cultivar
Oryzica Llanos 5 (Tohme et al., 1992).
Decreases in the use of fungicides as a result of farmers growing these new
varieties have been reported. Unfortunately, it has not been possible to review these
data at this time. Measures of declining use of fungicides in agroecosystems of
Colombia can be estimated, in that farmers’ expenditures on these chemicals range
from 6 to 50% of total crop protection costs. Actual estimates of how much farmers
have saved over this period of time and how much the use of fungicides has been
reduced as a result of resistance will be obtained later (F. Correa-Victoria, personal
communication).
The Case of Bacillus thuringiensis and Transgenic Crops
By using genetic engineering, it is possible to introduce novel sources of insect

resistance to crop plants. In this case, resistance comes from genes encoding the
production of various endotoxins, which is being done by some of the international
programs as shown in Tables 2 and 3, including work on maize. Engineered varieties
would eventually be suitably adapted for growth in areas of Latin America, some
areas of which are associated with centers of diversity for maize. It is essential to
prepare Latin American countries for the advent of transgenic maize containing
genes for insect resistance, for which it is claimed that dependence on pesticides
would be eliminated, thereby enhancing the quality of the agroecosystem.
Studies on the introduction of transgenic maize in Mexico were one of the cases
selected by IBS for the Latin American seminar. Serratos (1998) stated that research
criteria for transgenic corn to be introduced in Mexico should be based on charac-
terization of agroecological, social, and economic aspects of the area where it is to
be grown. The introduction of transgenic cultivars seems inevitable to developing
countries. Thus, it is important to consider the impact of transgenic cultivars on the
agroecosystems of countries with extensive diversity of native germplasm.
Research on the use of endotoxins in maize is also being done on tropical maize
at CIMMYT’s (Centro Internacional de Mejoramiento de Maiz y Trigo) Applied
Biotechnology Center. These activities include screening of cloned B.t. genes for
toxicity against Heliothis zea and other tropical maize pests. They are also working
on the transformation of tropical maize inbreds containing cry gene constructs and

greenhouse evaluations of acquired transgenic germplasm containing cry gene(s)
and introgression of cry gene(s) into tropical germplasm (IBS, 1994).
Research at CIMMYT and by commercial companies on hybrid maize suitable
for growth in tropical climates suggests the need for further study of their potential
effects on these complex agroecosystems. Thus, it is important to study, as a multi-
institutional task, gene flow and biological risks which may be associated with
transgenic maize in Mexico (Serratos, 1998). This could include genetic flux, hybrid-
ization, and introgression among the transgenic cultivators, native cultivators, and
wild parents. Addressing factors such as these would contribute to an analysis of
benefits from the transgenic maize in relation to potential environmental concerns.
In the final case (Whalon and Norris, 1998), the role of resistance management
when deploying transgenic B.t. plants is discussed within a resistance management
framework. Here, it was noted first that transgenic technology will help reduce
reliance on chemicals, reduce environmental contamination, and reduce human health
impacts by conventional pesticides. Second, this technology appeals to developing
countries lacking effective pesticide safety regulations because transgenic plants do
not carry the human and environmental risks that conventional pesticides do.
However, it was argued that some type of management is needed to sustain the
effectiveness of these pest control tactics by managing the factors that may contribute
to resistance development. This may require commitment and participation by farm-
ers, pesticide or seed suppliers, and regulators to help prevent insect resistance
through detection and proactive management. The preservation and management of
genetic resources, i.e., susceptible genes, is the key goal of resistance management
(Whalon and Norris, 1998).
The authors recommend that, as regards a specific group of technologies, the
decision to deploy transgenic crop plants should be based on an assessment of
indigenous ecological, environmental, socioeconomic, and agricultural conditions.
Criteria to consider include the risk of gene transfer from transgenic plants to related
species, availability of refugia to counteract resistance development, economic
importance of the target pests, and the level of cooperation among growers and
industry in the management of transgenic resources. The assessment should include
input from scientists, policy makers, agricultural specialists, public and private
institutions, and local farmers.
QUALITY INDICATORS AND NEW TECHNOLOGIES —

SYNTHESIS OF ABOVE DISCUSSION
Concerns regarding the use of crops modified by new technologies vary, as shown
by the case of rice and for B.t. technologies. Clearly, more issues are expected for
the use of products containing B.t.-derived genes. These differences point to the
need for some of the international crop biotechnology programs (see Table 2) to
consider their research, testing, and use of products in the context of integrated pest
or resistance management can be anticipated. It may also require more-detailed
consideration of the two indicators of agroecosystem quality presented in the section
on Quality Indicators — Linking Biodiversity with New Technologies, above.

The need for such approaches is often discussed in reports and workshops
enumerating biosafety considerations for the introduction of transgenes into tropical
agroecosystems. By summarizing these reports (see World Bank, 1993; Frederick
et al., 1995; Frederiksen et al., 1995; Beachy et al., 1997; Hruska and Pavon, 1997;
Serratos 1998; Whalon and Norris, 1998), the more specific considerations regarding
biosafety can be covered by the following categories:
• Transgene flow in centers of diversity: crops becoming weeds, transgenes moving

to wild plants, or erosion of genetic diversity
• Development of new viruses
• Resistance developed rapidly to the transgenes
• Affects on unintended targets
• Other ecosystem damage
Addressing these concerns begins with technical solutions, including data col-
lection and experimentation. However, there is also a more adaptive component
found in biosafety considerations, indicating agroecosystem complexities, the stake-
holders involved, and the need for information addressing the two quality indicators
selected. Generally, the more adaptive components of these concerns are voiced in
terms of educating policy makers and public regarding consequences of use and
deregulation, developing educational materials, and providing cost/benefit analysis
reflecting the needs or priorities of each country. These points are often raised by
participating countries during IBS policy seminars, and at biosafety meetings where
this topic is stretched to accommodate other debates. These more adaptive challenges
relate directly to the policy and management challenges facing leaders in developing
countries seeking to employ the products of new agricultural technologies.
Initiating programs to address some of the above considerations often exceeds
the funding base provided for the international programs. However, some of the
international programs have begun this experimentation and data collection, as is
being done for rice (Gould, 1997). There is an equally great need to build such
understanding among those responsible for agricultural research in the developing
countries. Unfortunately, developing countries cannot derive much information from
analysis by regulatory agencies in developed countries for permits or notification
for small-scale field testing of transgenic products, because the trial is conducted
within parameters taking into account isolation, pollen flow, and avoiding persistence
of crops at field sites.
These criteria and parameters enable those conducting tests to demonstrate that
transgenic plants are as safe as other plant varieties. However, such isolation practices
established for the needs of trials in the U.S. and Europe do little to satisfy the
concerns (as listed above) anticipated for tropical ecosystems or centers of diversity.
Of course, this is not the purpose of trials carried out in developed countries. The
questions are: who will determine and how, whether the new plants are of no greater
danger to tropical ecosystems than plants produced traditionally, and how will
technical estimates for the two quality indicators be prepared in this regard?

AGROECOSYSTEM QUALITY AND CHALLENGES AHEAD —
ADAPTIVE PROBLEMS REVISITED
The various points to be covered in this chapter are now complete, as summarized
in Table 1. While it is not common to pose agricultural questions in the context of
technical and adaptive problems, this distinction has much to offer discussions
concerning biotechnology, especially when considering the range of questions that
may be asked by various stakeholders regarding agronomic inputs and biodiversity.
For biotechnology-derived improvements to have acceptability, clear demonstrations
of utility with regard to stakeholder concerns for environmental and productivity
considerations are needed.
As mentioned above, agroecosystem quality may be improved by eliminating
or minimizing dependence on chemical inputs (quality indicator 2), although clear
data on this is lacking at the present time. They may also affect perceptions
regarding biodiversity (quality indicator 1) leading to widespread use of a variety
or, in the case of transgenic maize, have implications for gene transfer in a center
of diversity, or on horizontal gene transfer (Harding, 1996). The examples used
(durable blast resistance and B.t. technologies) indicate potential suitability for
farmers lacking access or money for chemical inputs, where it is desired to reduce
chemical inputs in traditional systems or where minimal disruption of biological
populations is desired. In the case of tropical maize with insect resistance, since
the technologies have not yet been used or tested in the field, it was not possible
to obtain estimates on expected decreases in the use of pesticides, as related to the
second quality indicator.
As seen in the policy seminars, new products often focus attention on acceptance
issues, which can be related to indications of agroecosystem quality. Consequently,
in each seminar, socioeconomic methodologies are explored in regard to how stake-
holders benefit from investments in biotechnology, and how such analysis can con-
tribute to the learning required to address environmental and productivity questions.
Follow-up to the seminars gives attention to identified needs, providing the oppor-
tunity to approach them as adaptive problems, often requiring changes in stakeholder
values, attitudes, or behavior.
This supports points emphasized by Whalon and Norris (1998), as much remains
to be learned regarding the wise management and deployment of genes introduced
through biotechnology. Thus, findings point to where future work can be anticipated
that it is hoped will diminish the learning required for adaptive situations. In many
cases, these situations will weigh productivity issues of profitability, market accept-
ability, and overall agronomic performance with effects on agroecosystem quality.
Neither dimension (environment or productivity) can be ignored. At the present
time, adaptive problems arising from international biotechnology efforts are
encountered not in the context of agroecosystem quality, but under the heading of
biosafety considerations. The relation among biosafety, solutions offered by bio-
technology, and more complex considerations of ecosystem effects is seen at many
workshops.

New biotechnologies used by farmers will raise adaptive problems, of which
biosafety deliberations may be one part. Stakeholder involvement will be essential
in considering these cases, especially given that local land-use knowledge continues
to be essential to food production in the tropics and in traditional agroecosystems
(Gliessman, 1993). Such knowledge reflects experience gained over many genera-
tions, and can contribute much toward sound management practices. Using local
knowledge when determining quality indicators could be done in conjunction with
efforts to determine natural resource or ecologically sound management practices.
However, as already stated, such measurements have human biases or judgments
attached to them and reflect the stakeholders involved.
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Biodiversity in Agroecosystems (Advances in Agroecology) - CRC Press (1998)

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BIODIVERSITY

Wanda W. Collins and

© 1999 by CRC Press LLC.

Library of Congress Cataloging-in-Publication Data

Biodiversity in agroecosystems / edited by Wanda Williams Collins, Calvin O. Qualset.

© 1999 by CRC Press LLC.

No claim to original U.S. Government works

© 1999 by CRC Press LLC.

© 1999 by CRC Press LLC.

© 1999 by CRC Press LLC.

Calvin O. Qualset, Ph.D., is Director of the Genetic Resources Conservation

© 1999 by CRC Press LLC.

Stuart S. Bamforth Wanda W. Collins (Editor)

© 1999 by CRC Press LLC.

Thomas E. Lacher, Jr. Keith S. Pike

© 1999 by CRC Press LLC.

© 1999 by CRC Press LLC.

© 1999 by CRC Press LLC.

© 1999 by CRC Press LLC.

Agroecosystems comprise 30% of Earth’s surface (Altieri, 1991). Microbial

© 1999 by CRC Press LLC.

species with little attention being given to microorganisms. Without microorganisms

© 1999 by CRC Press LLC.

The number of known microbial species is estimated to be over 110,000, but

© 1999 by CRC Press LLC.

SOIL MICROORGANISMS IN AGROECOSYSTEMS

The soil is full of microorganisms such as bacteria, actinomycetes, fungi, algae,

© 1999 by CRC Press LLC.

Plants may be an important selective force for the diversity of rhizosphere

© 1999 by CRC Press LLC.

MICROBIAL IMPACTS ON AGROECOSYSTEMS

Table 2 Several Functions of Soil Microbes

© 1999 by CRC Press LLC.

© 1999 by CRC Press LLC.

Soil microorganisms constitute a large dynamic source and sink of nutrients in

© 1999 by CRC Press LLC.

AGROECOSYSTEM IMPACTS ON MICROORGANISMS

Microbial diversity considerations need to be included in soil quality investiga-

Microbial communities in agroecosystems change with management history. In

© 1999 by CRC Press LLC.

POTENTIAL APPLICATION OF MICROBIAL INDICATORS

© 1999 by CRC Press LLC.

The diversity of microorganisms is greater than any other group of organisms

Support from the U.S. Department of Agriculture, Agricultural Research Service

© 1999 by CRC Press LLC.

© 1999 by CRC Press LLC.

© 1999 by CRC Press LLC.

© 1999 by CRC Press LLC.

© 1999 by CRC Press LLC.

© 1999 by CRC Press LLC.

© 1999 by CRC Press LLC.

Soil Microfauna: Diversity and

Agricultural plant production depends upon the decomposition of plant and

ROLE OF SOIL PROTOZOA

© 1999 by CRC Press LLC.

MEASURING PROTOZOAN BIODIVERSITY

© 1999 by CRC Press LLC.

PROTOZOAN DIVERSITY IN AGROECOSYSTEMS

Studies on grasslands (McNaughton, 1977; Tilman, 1996) show that biodiversity