Term: August 1999 through July 2001
Sponsor: Professor Robert Evenson
Economic Growth Center
Research: Agricultural development,
economic growth, and sustainable resource use
the past year, several research projects have come
to fruition. Most notably, Dr. Gollin expects to
complete a book manuscript edited jointly with Yale
Economics Professor Robert E. Evenson in January
2001. This manuscript summarizes the results
of a two-year effort by a group of economists from
around the world to examine the impact of
international agricultural research on the
sustainable production of crops. Their
findings have been presented at the World Bank and
in numerous other forums, and it is anticipated that
the book itself will receive considerable attention
within the international community. A major
finding of this research is the overwhelming success
of international research in developing improved
crop varieties over the past 40 years. These
varieties have played a substantial and important
role in contributing to unprecedented increases in
per capita availability of food. They have
also helped to alleviate stress on land and forest
resources by allowing for increased human
populations to be fed at least partly through
intensification of agriculture on existing crop
land, rather than through clearing of new land.
believed that this work will play a significant role
in shaping attitudes within the international policy
community. The study was commissioned by the
Technical Advisory Committee of the Consultative
Group on International Agricultural Research (CGIAR),
the umbrella organization that supports some 16
international agricultural research centers.
The study comes at a time when donors have been
reluctant to allocate funds for further agricultural
research, at least in part based on some
misconceptions about the continued impacts of such
on this research, Professor Evenson and Dr. Gollin
are organizing a session at this year’s annual
meetings of the American Association for the
Advancement of Science (AAAS). This session
will address the appropriate role for the public
sector in the further development of biotechnology
for agriculture. In addition to organizing the
session, they are also preparing one of the papers,
which Dr. Gollin will present. The main
message of the paper is that public sector research
will remain the predominant source of technical
change for agriculture in developing countries.
For most major crops, we see little evidence that
the private sector will carry out much research
relevant to the needs of poor countries.
accomplishment during the past year was the
publication of a paper on the management of
materials in agricultural gene banks. For most
cultivated crops, genetic
diversity is collected and conserved ex situ in gene
banks – typically cold storage facilities designed
to maintain the viability of seeds for 50 to 100
years. Along with colleagues from CIMMYT, the
International Center for Maize and Wheat Improvement
in Mexico, Dr. Gollin wrote a paper that addressed
some practical questions relating to the use of gene
banks for breeding improved varieties. This
paper, recently published in the American Journal of
Agricultural Economics, has been well received by
plant breeders and gene bank managers, as well as by
Gollin has also continued to explore theoretical
models of the relationship between agricultural
development and economic growth. Along with
several colleagues at Yale and elsewhere, he is
working on papers that focus on the theoretical and
empirical relationships between technical change in
agriculture and economic growth.
of his research, he has taken advantage of many
intellectual resources at Yale and has enjoyed the
chance to interact with students and faculty in the
Economics Department, the School of Forestry and
Environmental Studies, and the Yale Center for
International and Area Studies. His experience
at Yale has been both productive and enjoyable and
he thanks the Donnelley family and the Yale
Institute for Biospheric Studies for their kind and
Term: September 1999 through August 2001
Sponsor: Professor Oswald Schmitz
School of Forestry and Environmental Studies
Research: Testing the effect of state dependent
decision-making of individual herbivores on food web
the origin and maintenance of global patterns of
biodiversity is one of the monumental problems of
biology (Ricklefs and Schluter 1993).
Ecological studies of the past thirty years tended
to focus on population level processes in order to
explain community structure. However, this
approach failed to explain global biodiversity
patterns (Ricklefs and Schluter 1993). These
studies as many others at the population and
community levels were based only on numerical
effects and did not take into account the possible
behavioral effects. Recently, it has been
shown that individual behavior may stabilize
population dynamic (Mangel and Roitberg 1992), drive
trophic cascades (Schmitz et. al. 1997), and be
critical to persistence of populations in the
landscape (Lima and Zollner 1996; Roitberg and
Mangel 1997). These studies as some others
have led ecologists to the conclusion that one
should not ignore the role of individual behavior
when studying the structure and the function of
complex ecological entities (e.g., Sutherland 1996;
Fryxell and Lundberg 1998).
Ovadia’s research aim is to incorporate state
dependent decision making by individual herbivores
into a model of food web dynamics, and to test this
model experimentally in the field.
Ovadia uses the Gecko (Booth 1997; Schmitz and Booth
1997; Schmitz 2000) as a tool for studying the
effect of state dependent decision-making by
individual herbivores on food web dynamics.
The Gecko is a spatially explicit individual based
model developed at the CCE (Center of Computational
Ecology) to explore the effect of individual
behavior on food web dynamic (Booth 1997; Schmitz
and Booth 1997; Schmitz 2000). He has modified
the Gecko to include state dependent foraging
decisions by individual herbivores and ran
simulations and used the results to design and to
generate predictions for the field study.
study is located in an old-field located at the
Yale-Myers Research Forest in northeastern
Connecticut. The food web includes three-trophic
levels, spider predators, grasshopper herbivores and
field study is divided into two major parts:
behavioral experiments and food web experiments.
During the behavioral experiments Dr. Ovadia
monitors the foraging behavior of grasshoppers with
different internal Body State exposed to different
levels of predation risk. During the food web
experiments, he tests the effect of different levels
of chronic internal Body State of grasshoppers on
1999 – April 2000:
Theoretical study which includes three parts: 1)
Modifying the Gecko to include state dependent
decision making by herbivores, 2) Running
simulations to test the effect of herbivores state
on food web dynamic, and 3) Using the simulations
results to design and to generate predictions for
the field study.
2000 – September 2000:
Field study that includes two major parts: 1)
Behavioral experiments, and 2) Food web experiments.
2000 – April 2001:
Analyzing the results from the empirical study.
Using the first year results to enhance the
development of the theoretical study.
2001 – September 2001:
Field study testing the predictions generated by the
Gecko during the theoretical study of the second
progress done since Dr. Ovadia’s arrival at the
Dr. Ovadia is working in collaboration with Ginger
Booth from the CCE. They have modified the
Gecko to include state dependent foraging decision
by the herbivores in the model. He ran
simulations and used the results to design and to
generate predictions for the field study.
Dr. Ovadia did a large field experiment during the
summer and is currently analyzing the data.
Dr. Ovadia is working on his PhD publications and
has submitted two papers, one accepted for
publication in Behavioral Ecology and the other
currently is in review.
Dr. Ovadia is working on two papers: the first
manuscript is a theoretical state dependent Gecko
paper and the second one is an empirical paper that
is based on the results of field study.
Booth, G. 1997. Gecko: A continuous 2D
world for ecological modeling. Artificial
Life. 3: 147-163.
Fryxell, J.M. and Lundberg, P. 1998.
Individual Behavior and Community Dynamics.
Chapman and Hall, London.
Lima, S.L. and Zollner, P.A. 1996.
Towards a behavioral ecology of ecological
landscapes. Trends in Ecology and Evolution.
Mangel, M. and Roitberg, B.D. 1992.
Behavioral stabilization of host-parasite population
dynamics. Theoretical Population Biology.
Ricklefs, R.E. and Schluter, D. 1993.
Species Diversity in Ecological Communities,
Historical and Geographical Perspectives. The
University of Chicago Press, Chicago.
Roitberg, B.D. and Mangel, M. 1997.
Individuals on the landscape: behavior can mitigate
landscape differences among habitats. Oikos.
Schmitz, O.J. 2000. Combining field
experiments and individual-based modeling to
identify the dynamically-relevant organizational
scale in a field system. Oikos. In
Schmitz, O.J., Beckerman, A.P. and O’Brien, K.M.
1997. Behaviorally mediated trophic cascades:
effects of predation risk on food web interactions.
Schmitz, O.J. and Booth, G. 1997.
Modeling food web complexity: The consequences of
individual-based, spatially explicit behavioural
ecology on trophic interactions. Evolutionary
Ecology. 11: 379-398.
Sutherland, W.J. 1996. From Individual
Behaviour to Population Ecology. Oxford
University Press. Oxford.
Term: July 2000 through June 2002
Sponsors: Dr. Gisella Caccone and Professor
ECOSAVE Conservation Genetics Laboratory
Research: “Research in Molecular
Claudio Ciofi arrived at Yale on July 1, 2000 as a
Gaylord Donnelley Environmental Fellow. Dr.
Ciofi received his Ph.D. in Conservation Biology
from the University of Kent in England in 1998.
His dissertation was on the spectacular Komodo
dragon and is one of the finest studies done on the
Komodo dragon in the last 20 years. He
conducted arduous fieldwork in an effort to
understand these lizards, including tagging them and
determining range size. He also collected
blood from about 150 animals and conducted
microsatellite DNA studies. Finally, he
analyzed the genetic data using sophisticated
techniques that allowed him, among other things, to
draw conclusions about the relative isolation among
the remaining populations. Thus he has
demonstrated all the skills required in modern
conservation biology: fieldwork, ecological studies,
laboratory genetic studies, and sophisticated
computer-based analysis. In addition, he has
been working with and training local Indonesians,
not only to give them skills, but also to raise
their appreciation for these endangered animals with
the hopes of affecting public policy.
his arrival at Yale University as a Donnelley
Post-Doctoral Fellow, he has been involved in a
number of very productive research and educational
activities, both at Yale and abroad, all part of the
two main wildlife conservation projects that he has
been involved in. The giant Galápagos
tortoise and the Komodo dragon projects are
well-established studies where molecular genetic
techniques are employed to better define the
distinctiveness of natural populations, to obtain
information on demographic parameters, and to
provide recommendations on the amount of effort that
should be devoted to conservation. This is a
fast evolving discipline, which is being integrated
with ecological and
demographic studies, to formulate appropriate
management strategies for endangered species.
the first three months of his appointment, he has
participated in scientific expeditions in Ecuador
and Indonesia to collect samples for genetic
analysis from giant tortoises and Komodo dragons
respectively. The Ecuadorian expedition was
carried out in the Galápagos archipelago, about
1,500 km off the coast of Ecuador in the Pacific
Ocean. The fieldwork was conducted on Isabela,
the largest and geologically youngest of a set of 15
volcanic islands. The study, organized at
Yale, included four researchers of different
nationalities. Its purpose was to collect
blood samples of giant tortoises inhabiting the
slopes of two volcanoes on the north part of the
island. The group identified, measured, sexed,
and sampled 93 tortoises of up to 200 kg in weight.
Samples were collected from the brachial vein
located on one of the forelimbs. During the
work, Dr. Ciofi had the opportunity of training two
Ecuadorian undergraduate students in the collection
and storage of reptile blood.
Indonesian expedition was conducted at Komodo
National Park in the southeast part of the country.
The goal of the study was to collect blood samples
from adult females once the location of their nests
was identified. Subsequent collection of
samples from the hatchlings will allow a DNA
fingerprinting analysis to assess single or multiple
paternity. This will help to clarify the
mating system of the species. Dr. Ciofi
collected blood samples from five females.
As for his previous studies on the Komodo dragon,
specimens were caught in baited traps (300x50x50
cm). Young females were first restrained, and
500 micro liters of blood were collected from the
caudal vein using a 5 ml syringe and a spinal needle
(0.70x90 mm), while large specimens were kept in the
trap and bled by nail-clipping.
genomic DNA was extracted from blood samples soon
after fieldwork. Genetic analysis is now
conducted by assessing allelic differences among
individuals at nine microsatellite loci.
Microsatellites are short, tandemly repeated simple
sequences of nuclear DNA. Genetic differences
between individuals are determined by identifying
variation in the number of repeats.
Microsatellite loci are first amplified by
polymerase chain reaction (a technique to obtain
sizable specific DNA sequences from minute amounts
of whole DNA), and individual genotypes are then
obtained. Differences in allele sizes between
individuals as little as one DNA base pair can be
detected, allowing the maximum degree of resolution.
Yale undergraduate students are currently working on
the above techniques under Dr. Ciofi’s
supervision. During their training period,
they will become proficient in DNA extraction and
analysis methods and will gain knowledge of how the
data obtained can be used to design and implement
wildlife management plans.
Komodo dragon study is part of a wider project Dr.
Ciofi set up during his Ph.D. work. At
Yale, he has had the opportunity to continue and
expand the work he initiated using genetics to
obtain quantitative information on population
divergence, reproductive biology, and demography of
addition, he has submitted a number of applications
for funding to US Institutions to support his
ongoing research. In January 2001, three
grants have been awarded by private Zoological
foundations, and additional support has been
confirmed by the Smithsonian Institution, Washington
D.C. These funds will allow Dr. Ciofi to
continue his studies in the laboratory and in the
field, and will also afford him the opportunity to
continue his collaboration with Indonesian
universities and support undergraduate short-term
field studies on the biology and management of the
Ciofi is also collaborating with the Zoological
Society of San Diego and they have begun to fund a
capacity building project at the University of Bali
and Komodo National Park. He produced a
memorandum of understanding with the Indonesian
Department of Nature Protection and Conservation,
Udayana University (Bali, Indonesia) and the
Zoological Society of San Diego, to set up a
molecular biology laboratory in the department of
Basic Science at Udayana, and a field research
station in Komodo National Park. In 2001 Dr.
Ciofi will be directly involved in setting up the
administrative, logistic and scientific part of this
initiative. Funds will be provided for
capacity building, one Post-Doctoral position, and
stipends for a local member staff person, and
scholarships for Indonesian graduate students.
Term: September 2000 through August 2002
Sponsors: Professor Mark Ashton,
School of Forestry and Environmental Studies
Professor Michael Donoghue,
Dept. of Ecology and Evolutionary Biology
Research: “Study of the Phylogenetic
Structure of Forest Tree Communities in Indonesian
Borneo and at Selected Comparative Sites in Asia and
Campbell Webb arrived at Yale on September 1, 2000
as a Gaylord Donnelley Environmental Fellow.
He received his Ph.D. from Dartmouth College in
Ecology and Evolutionary Biology and studied the
regeneration ecology of the tree community at the
Gunung Palung National Park, Kalimantan, Indonesia,
and its role in the maintenance of species diversity
for his doctoral work. In a study of 150
species, he found that seedlings of common species
generally suffer high mortality than those of rare
species, an exciting discovery
that implies the existence of a ‘balance of
nature’ in these economically valuable forests.
arriving at Yale, Dr. Webb spent the last two years
as a research fellow of the Arnold Arboretum,
Harvard University, working on regional-scale
patterns of tree species diversity in Borneo and
developing methods to investigate the phylogenetic
structure of sympartric plant communities. He
also gained experience in ‘alpha-taxonomy’,
describing several new plant species from Borneo.
research explores the origin and maintenance of tree
species diversity in tropical forests. He has
assessed a number of hypotheses for the contemporary
coexistence of tree species, and found evidence that
there exists a `balance of nature' in Bornean
forests, in that regeneration of common species is
less successful than that of rare species.
Recently, he has started to examine the potential
for modern phylogenetic analyses to be used in our
investigation of ecological community structure.
Since he has come to Yale, Michael Donoghue and he
have begun to develop the tools and theories needed
for this new approach. Mark Ashton was
on sabbatical for most of the Fall of 2000, and Dr.
Webb looks forward to working with him in 2001.
his primary research focus is on the theory of
forest community structure, he has long been active
in rain forest conservation, particularly in the
inventory and mapping of threatened forest areas.
As described below, he has made progress in these
areas. The general program of seminars at the
Yale School of Forestry and Environmental Studies
has been a great resource. The country that he
has been most involved in is Indonesia, and a
pleasant surprise for him at Yale has been the
existence of a sizable body of Indonesian graduate
students, many of who work on environmental issues
he finds Yale to be an excellent home for his
research and conservation activities, and he
expresses his deep gratitude to the Yale Institute
for Biospheric Studies, and especially to the
Donnelley family for supporting his fellowship.
July - December 2000
Any analysis of the phylogenetic structure of
communities will depend upon having phylogenies for
the taxa in the communities. However, very few
of these taxa will have ever been the subject of
detailed molecular analysis. We must depend
therefore on constructing `supertrees' of the
communities, using published phylogenies of closely
related species. Dr. Webb has developed a
system for assembling these supertrees that includes
a frequently-updated `backbone' phylogeny for the
higher plants. See the project website at http://pantheon.yale.edu/~cw252/ttp.
The analyses of community phylogenetic structure
that is planned depends upon specialized software.
This software is based on code that Dr. Webb wrote
before arriving at Yale, but it must be continually
modified and upgraded. The package of software
now includes measures of the tree-balance of
Professor Michael Donoghue and Dr. Webb were awarded
a grant from the Biotic Surveys and Inventory panel
of NSF to develop methods for biodiversity surveys
using wireless technology.
Professor Michael Donoghue and Dr. Webb also
submitted a proposal to Annual Review of Ecology and
Systematics for a review paper entitled
``Phylogenies and Community Ecology,'' to be written
with David Ackerly, Mark McPeek and Michael Donoghue.
The proposal was successful and the paper is due
proposal was also submitted to the National Center
for Ecological Analysis and Synthesis to organize a
15-person working group, also under the title,
``Phylogenies and Community Ecology.''
Dr. Webb attended the Ecological Society of America
meetings in Snowbird, Utah. Discussed wireless
biology techniques at a special session of the
Long-Term Ecological Research meeting and presented
a talk entitled, “Seed dispersal determines local
seedling species diversity in Bornean rain forest”
at the General Annual Meeting.
Dr. Webb visited Steve Hubbell at University of
Georgia to collaborate on analysis of the
phylogenetic structure of the 50-ha forest research
plot on Barro Colorado Island, Panama.
Dr. Webb visited David Ackerly and Miguel
Martinez-Ramos at Stanford University, to begin
collaboration on a simulation model of community
assembly and phylogenetic structure.
the NSF grant (above), Dr. Webb has started to
assemble a wireless system that links a field
collector with the global systematics and education
community. This will enable the collector to
more efficiently inventory taxa outside their area
of expertise, and will also be a powerful tool for
engaging students in biodiversity studies. The
current setup comprises a PalmPilot with PalmPix
camera connected to a laptop computer up to 20 miles
away, via a FreeWave spread-spectrum radio.
The laptop will eventually be positioned at a
base-camp, and will transmit data to a web-server,
via a cell- or satellite telephone. The
web-server will post images and messages on the
Internet, and route replies back to the user.
He will test the setup in Borneo during a field trip
in March and April 2001.
Computer-based identification keys provide a
powerful tool for anyone trying to identify unknown
taxa. However, even laptops are too cumbersome
to carry to the field, where identification is most
needed. However, ‘palm-tops’ are small
enough, and Dr. Webb developed a simple
identification application for the PalmPilot, called
‘PalmKey.’ The tool can be downloaded from
his website: www.herbaria.harvard.edu/~cwebb/PalmKey.
Planned for 2001
Continued work on phylogenetic structure of forest
communities: compiling datasets, developing
software, publishing findings.
NCEAS meetings (2 x) for working group entitled:
“Life-history variation and community structure in
neotropical rainforest communities: ecological and
Possible NCEAS meeting for working group entitled
“Phylogenies and community ecology.”
Ecological Society of America meeting, Madison WI.
Invited symposium speaker.
Association of Tropical Biology meeting, Bangalore,