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YIBS 1999-2000 Annual Report

III.  GAYLORD DONNELLEY ENVIRONMENTAL FELLOWSHIP

Donnelley Environmental Fellows Appointed 1999 - 2000

Dr. Douglas Gollin
Term:  August 1999 through July 2001
Sponsor:  Professor Robert Evenson
Economic Growth Center
Research:  Agricultural development, economic growth, and sustainable resource use

Over 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.

It is 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 work.

Drawing 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.

Another 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 economists.

Dr. 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.

Outside 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 generous support.

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Dr. Ofer Ovadia
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 dynamics.

Scientific Background:

Understanding 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).

Dr. 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.

The Model:

Dr. 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.

The Empirical Study:

The 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 herbaceous plants.

The 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 community dynamic. 

Research Plan:

September 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. 

April 2000 – September 2000:
Field study that includes two major parts: 1) Behavioral experiments, and 2) Food web experiments.

September 2000 – April 2001:
Analyzing the results from the empirical study.  Using the first year results to enhance the development of the theoretical study.

April 2001 – September 2001:
Field study testing the predictions generated by the Gecko during the theoretical study of the second year.

The progress done since Dr. Ovadia’s arrival at the Yale University:

1)      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.

2)      Dr. Ovadia did a large field experiment during the summer and is currently analyzing the data.

3)      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.

4)      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.

References:

1)  Booth, G.  1997.  Gecko: A continuous 2D world for ecological modeling.  Artificial Life. 3: 147-163.

2)  Fryxell, J.M. and Lundberg, P.  1998.  Individual Behavior and Community Dynamics.  Chapman and Hall, London.

3)  Lima, S.L. and Zollner, P.A.  1996.  Towards a behavioral ecology of ecological landscapes. Trends in Ecology and Evolution.  11(3):131-135.

4)  Mangel, M. and Roitberg, B.D.  1992.  Behavioral stabilization of host-parasite population dynamics.  Theoretical Population Biology.  42:308-320.

5)  Ricklefs, R.E. and Schluter, D.  1993.  Species Diversity in Ecological Communities, Historical and Geographical Perspectives.  The University of Chicago Press, Chicago.

6)  Roitberg, B.D. and Mangel, M.  1997.  Individuals on the landscape: behavior can mitigate landscape differences among habitats.  Oikos. 80:234-240.

7)  Schmitz, O.J.  2000.  Combining field experiments and individual-based modeling to identify the dynamically-relevant organizational scale in a field system.  Oikos.  In Press.

8)  Schmitz, O.J., Beckerman, A.P. and O’Brien, K.M.  1997.  Behaviorally mediated trophic cascades: effects of predation risk on food web interactions.  Ecology.  78(5):1388-1399.

9)  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.

10)  Sutherland, W.J.  1996.  From Individual Behaviour to Population Ecology.  Oxford University Press.  Oxford.

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Donnelley Fellows Appointed for 2000 – 2002

Dr. Claudio Ciofi
Term:  July 2000 through June 2002
Sponsors:  Dr. Gisella Caccone and Professor Jeffrey Powell
ECOSAVE Conservation Genetics Laboratory
Research:  “Research in Molecular Ecology”

Dr. 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. 

After 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.

In 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.

The 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.

Whole 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.

The 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 the species.

In 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 Komodo dragon.

Dr. 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.

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Dr. Campbell Webb
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 Central America”

Dr. 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.

Before 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.

His 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.

While 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 back home. 

Overall, 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.

Activities, July - December 2000

Main Research

1) 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.

2) 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 community phylogenies.

Proposals and Grants

1) 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.

2) 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 January 2002.

3) A 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.''

Visits and Meetings

1) 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.

2) 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.

3) 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.

Biological Inventory

1) With 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.

2) 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.

Activities Planned for 2001

1) Continued work on phylogenetic structure of forest communities: compiling datasets, developing software, publishing findings.

2) NCEAS meetings (2 x) for working group entitled: “Life-history variation and community structure in neotropical rainforest communities: ecological and phylogenetic influences.”

3) Possible NCEAS meeting for working group entitled “Phylogenies and community ecology.”

4) Ecological Society of America meeting, Madison WI.  Invited symposium speaker.

5) Association of Tropical Biology meeting, Bangalore, India.

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YALE INSTITUTE FOR BIOSPHERIC STUDIES
Oswald Schmitz, Director
Rose Rita Riccitelli, Assistant Director
LaToya Sealy, Sr. Administrative Assistant
Environmental Science Center, Room 132
21 Sachem St., P.O. Box 208105
New Haven, CT 06520-8105
Phone: (203) 432-9856 · Fax: (203) 432-9927