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Home , Evolutionary biology, Evolutionary physiology

Frontiers in Evolutionary

REPORT OF A WORKSHOP

PREPARED FOR

THE NATIONAL SCIENCE FOUNDATION

March 2005

Table of contents

Introduction ...... 1 Emerging tools ...... 1 Research themes ...... 2 The of genome structure and ...... 3 /evolution of ...... 3 divergence and ...... 4 of development ...... 4 Evolution of integrated ...... 5 Microbial systems ...... 6 Evolution of conflict and cooperation ...... 6 Large-scale patterns of diversity in time and space ...... 7 Applied evolution ...... 7 Institutional resources ...... 8 Infrastructural needs and opportunities ...... 9 : Workshop participants ...... 10

INTRODUCTION frontiers of evolutionary research in the coming Support for ecological and evolutionary research decade. As a result, the themes for future at the National Science Foundation expanded research outlined here reflect both new questions substantially during the past decade. To help and new conceptual frameworks, as well as the guide this expansion, in October 1998 the application of new tools to classic questions in program at NSF hosted a . workshop on “Frontiers in Population Biology”. The workshop had four specific goals: The report from that workshop identified and 1) To identify emerging tools essential to illustrated seven research themes of special evolutionary research; interest in population biology, and described how 2) To identify and illustrate research these themes connect to genetics/ on themes of particular promise; the one hand and to environmental issues on the 3) To summarize major institutional other. The report has been valuable to officers at resources available to support NSF in shaping their programs and balancing evolutionary research; their grant portfolios. 4) To suggest infrastructural needs and Our understanding of evolutionary biology opportunities for enabling the next has advanced enormously in the past six years. generation of advances in our understanding The explosion of genomic information, rapid of evolution. advances in computer and sensor technology, and the development of sophisticated new statistical and analytical approaches coupled with the EMERGING TOOLS recent reorganization of programs at NSF make Many research frontiers identified in this report this an appropriate time to evaluate progress and are now accessible only because of recent, to identify areas in which new investment is explosive advances in technology, while others warranted. In January 2005 a panel of experts lie at the fringe of current technology. A shared met at NSF to identify areas that represent the feature of many of these advances is the scale at frontiers for evolutionary research in the coming which they are conducted. We can now sequence decade. The scope of this workshop was whole genomes, analyze expression of all genes, somewhat larger than that considered at the 1998 and analyze data with desktop supercomputers. It workshop, reflecting the range of evolutionary is clear that advances in phenotypic analysis research and topics considered by Population and equivalent to those on the genomic and Evolutionary Processes, Ecological and computational side are needed, as these research Evolutionary , Animal and frontiers require a variety of high-throughput, related programs at NSF. In addition to high-precision techniques for measurement of identifying nine thematic areas within genetic, physiological, and phenotypic evolutionary biology, the panel considered how characteristics in large numbers of individuals in questions within these areas relate to questions in the field as well as in controlled, laboratory and genomics, , environments. environmental sciences and other areas. • Genomics – The advent of high throughput In reflecting on research frontiers that have DNA sequencing has led to the availability emerged since 1998, two related are of whole genome sequences for a broad evident. First, evolutionary approaches and phylogenetic sample of , including perspectives are increasingly an integral part of replicate genome sequences for different all areas of biological research, from molecular individuals of the same species. Microarray biology to macroecology. As a result, we are analyses of gene expression (and similar increasingly able to explore the mechanisms, methods) allow dissection of complex processes and patterns of evolutionary change at genetic networks involved in physiological, multiple levels of biological organization from developmental, and behavioral responses. the gene to the . Second, technological Large-scale genetic manipulation allows advances in genomics, computation, and experimental tests of adaptive hypotheses informatics have provided a world of new tools involving particular genes or sets of genes. and information to apply to evolutionary Environmental DNA provides insights into questions. These and similar technical advances the physiological and will continue to be a key factor in pushing the of unculturable microbes. Challenge: to

1 Figure 1: Connections among research frontiers identified in this report.

connect sequence-and expression-level data allow them to record corresponding with organismal phenotypes and responses. environmental variables at the scale at which • Analytics – The high-speed computational organisms sense the environment. Wireless resources now available enable complex technologies allow these measurements to statistical analyses and simulation studies be recorded without capturing or relocating faithfully reflecting details of biology that the individuals, and future integration with were previously impossible. Genome, global positioning systems will allow these phylogenetic, and environmental databases data to be integrated with fine-scale climate provide a wealth of information about the and environmental variables. Challenge: to genetic characteristics of organisms, their develop high-throughput methods allowing evolutionary relationships, and the external complex phenotypic measurements to be environment in which they occur. Continued made reproducibly at high frequency on a statistical and computational developments large number of individuals. are paving the way for a reconciliation of historical and cross-sectional analysis of . Challenge: to develop RESEARCH THEMES the conceptual framework for large-scale Research questions in evolutionary biology are databases of organismal phenotypes, the highly interrelated. Answering fundamental tools necessary to integrate these disparate questions about organismal adaptation, for databases into a comprehensive for example, requires understanding not only the analysis of large-scale patterns in genomic consistency, magnitude, and objects of natural and phenotypic evolution, and the statistical selection but the ways in which genome structure techniques for fine-scale evolutionary (including the structure of gene regulatory analysis of population history. networks) influences the distribution of mutational effects and the response to selection. • Phenomics – Advances in sensor technology enable real-time measurements Moreover, determining how genome structure of many important physiological variables in influences the evolution of integrated phenotypes the field. Advances in sensor design will is vital if we are to understand how individual-

2 level differences in development are related to • What are the extents and rates of changes in mechanisms of species divergence and large- genome structure and size, and what are scale patterns of biotic diversity. their functional consequences for organismal The workshop identified nine interrelated evolution? themes as research frontiers, broad areas where • How do new genes arise and go extinct? existing or foreseeable technology makes great • How does selection act on large-scale advances possible in the next decade (Figure 1). variation in genome structure and organization? The Evolution of Genome Structure and • What are the relative roles of changes in Function structural, regulatory, and non-genic Scope: Evolutionary change requires change in sequences in organismal evolution? the architecture or components of genomes, and • What evolutionary forces determine organismal properties determined by genetic or transposable element activity and number in molecular interaction networks that arise from genomes, and what are the functional the expression of genes. Understanding genomes, consequences of these mobile elements? their components, and the interaction networks • What is the structure of genetic networks, that arise from them is fundamental to a modern how do they evolve and how does network understanding of the genetic basis of influence organismal evolution? evolutionary change. • What are the evolutionary causes of Rationale: Genomes are highly complex differences in recombination and molecular assemblages that encompass the rates across the genome? fundamental physical organization of genetic information in organisms. Fundamentally, Genetics/evolution of adaptation genomes are organized to encode the genetic Scope: Adaptation requires both that organisms information in discrete units (genes) as well as to differ in and that those fitness differences contain sequence elements (e.g., centromeres and be heritable. Understanding how adaptive traits telomeres in most eukaryotic systems, ori evolve requires that we integrate genetic analyses sequences in bacteria) necessary for the of phenotypic differences with analyses of the transmission and maintenance of genetic performance consequences of phenotypic information. differences in an ecological context. Genomes also have specific architectures Rationale: Molecular genetic studies have and components that vary dynamically both provided a wealth of information on historical within and among species. We are beginning to patterns and rates of evolution of particular genes understand that single-gene approaches to but have rarely made a connection with understanding genome structure and function are particular phenotypes or . insufficient. Individual gene products are Similarly, a great deal is known about the embedded in large-scale interaction networks adaptive value of specific phenotypes in that represent integrated functional units at the ecological contexts and rates of phenotypic molecular genetic level. evolution, but little is known about their genetic Our understanding of the evolutionary basis. While connecting these types of studies dynamics of diversification in genome structures has been an underlying goal in evolutionary and their associated molecular genetic networks, biology since the synthesis of evolutionary however, remains limited. Moreover, few theory and genetics, only now has it become comparative genomic studies are underpinned by possible to make these connections explicit. New a clear phylogenetic context. Finally, we lack genomic, molecular and computational tools theoretical analyses to inform our understanding offer the opportunity to link genetic variation at of how genomes and their associated genetic the DNA level with phenotypes, even in non- networks evolve. The revolution in genomics model organisms. Combined with technological technologies and resources, including whole advances and novel theoretical machinery, we genome sequences, low-cost sequencing, now have an unparalleled ability to measure microarray technologies and computational selection on the genes underlying phenotypic power, now allow us to begin to address variation in natural . Furthermore, questions surrounding the evolution of genome improved understanding of gene functions and structure and function. interactions will allow us to study more fully the Frontier questions evolution of adaptive phenotypes. Investigations

3 into the existence of generalities concerning the architecture of traits that differentiate populations genetic basis of adaptive evolution will require and species. studies in many different organisms over long Frontier questions periods of time. • How do gene-genealogies diverge at Frontier questions speciation? • What is the molecular basis of adaptation? • What is the of • What types of genes are involved in behavioral, ecological, physiological and adaptation: regulatory or structural? developmental incompatibilities that cause • What types of are involved in reproductive isolation? How does genetic adaptation: non-coding, point, indel, architecture constrain or promote divergent transposable elements? selection on such traits? • What genetic features of adaptive evolution • Do pre-zygotic incompatibilities evolve are convergent? more frequently or less frequently than post- • To what extent is phenotypic convergence zygotic incompatibilities in the initial stages the result of genetic convergence? of divergence? • How can an understanding of the molecular • What proportion of the species’ genome is bases of adaptation reveal historical patterns impervious to ? What are the of adaptation in natural populations? consequences of porous species boundaries? What is the role of hybridization and Population divergence and speciation genome reorganization in speciation? Scope: Population divergence and speciation • Must gene flow be absent for speciation to are fundamental processes for the generation of occur? How common are the conditions biological diversity. They produce the boundaries under which can occur? within which microevolutionary forces act. They How common is reinforcement? are manifest by changes in the allelic • Is the standing genetic variation within composition of phenotypic variation among both species the major genetic reservoir for diverging populations within species and speciation? recently diverged species groups. • What can the patterns of variation among Rationale: New approaches and technologies taxa tell us about the evolutionary processes are facilitating unprecedented insights into the by which they arise? mechanisms of population divergence and speciation. The ability to apply these Microevolution of development technologies to non-model systems offers the Scope: All phenotypes in multicellular opportunity to probe a diversity of speciation organisms are the result of development. An patterns and mechanisms. This comparative emerging understanding of developmental frontier will enable syntheses of evolutionary mechanisms coupled with recently developed forces that cause phenotypic divergence and genomic tools makes it possible for the first time speciation, an understanding of genetic to study the evolutionary mechanisms by which architectures of diverging traits and populations, differences in organismal development arise. and predictions about the extent and direction of Rationale: The last twenty years of work in possible gene flow between species. Phenotypes developmental genetics has generated detailed of particular interest include those underlying information regarding shared and divergent prezygotic incompatibility, such as features of the developmental mechanisms that morphological and behavioral traits involving underlie phenotypic (primarily morphological) sexual or ecological isolation, and those differences among broad taxonomic groups. underlying postzygotic incompatibility, such as These studies have revealed the surprising result developmental or physiological traits causing that there is a shared set of developmental and sterility or inviability. Understanding how regulatory processes (a shared “developmental these traits evolve requires that we understand toolbox”) that serves as the foundation for the distribution of genetic variation within and development within plant and animal systems. among diverging species, the shape and How can we understand organismal diversity in of gene genealogies that give the face of this apparent pattern of conservation? historical insight into that genetic variation, Armed with a better grasp of large scale patterns genome and chromosomal structural variation of the evolution of development, it is now between recently formed species, and the genetic appropriate to turn our attention more fully to the

4 evolutionary processes that lead to these patterns. signaling and locomotory performance, but Rapid advances in the detailed analysis of understanding their evolution requires developmental mechanisms within model taxa approaches that treat the as an coupled with increasing availability of genomic integrated system rather than as a set of isolated and functional genomic information now allows traits. for detailed studies of variation in developmental Rationale: New approaches and technologies processes within populations. These studies have enabled to measure complex, include an analysis of molecular variation and integrated phenotypes and their functional of developmentally relevant consequences in increasing detail and loci, as well as variation in functionally related sophistication. An individual’s fitness is the characters such as levels and localization of gene result of an integrated suite of phenotypic traits expression. Complementary to the study of interacting with environmental conditions; and variation within populations is the examination many such traits are multi-dimensional functions of divergence in developmental characters of age and environmental conditions. However, among closely related species. The increasing evolutionary studies often characterize an availability of genomes of closely related taxa individual’s phenotype in terms of a small will facilitate this work. Another exciting area of number of phenotypic traits, and consider future research centers on placing this variation and selection of phenotypes in a simple, developmental information into an atomistic manner. Recent technological advances environmental and ecological context relevant to make it possible to pursue a different approach, the whole . to understand variation and selection of Frontier questions integrated phenotypes and to investigate how this • How does phenotypic variation arise from integration changes during evolution. An variation during development? How does important related challenge is to develop the selection act on this variation? capacity to measure multiple components of • What are the developmental mechanisms integrated phenotypes on the large number of underlying phenotypic differences among individuals required for lab and field studies of closely related species and how are these genetic variation, selection, and evolution. differences related to variation within- Frontier questions species? • What are the patterns of evolution in • What is the level of natural variation in integrated suites of morphological, genes affecting different stages of behavioral, physiological and other traits? development? What are the signatures of Are there different patterns or rates of selection and drift on these genes? evolution for different types of traits? • How do homologous developmental • How do we visualize and quantify patterns processes generate non-homologous of phenotypic and genetic variation in high- phenotypes (i.e., what is the developmental dimensional, integrated phenotypes? basis of convergence and parallelism)? • How modular are integrated phenotypes in a • How does the structure of developmental functional sense? How does functional pathways influence the evolution of the integration emerge during development? genetic components of those pathways? • How modular are integrated phenotypes • How is adaptive plasticity achieved through during evolution? How readily are such development modulation? What is the modules co-opted into new functional roles genetic architecture and molecular basis of during evolution? plasticity? • How do behavioral responses affect the pattern of selection on morphological, Evolution of Integrated Phenotypes physiological and other traits? How do Scope: Many diverse characteristics of an immune responses and behavior interact to organism—, physiology, behavior— influence their joint evolution? combine to yield an integrated phenotype that • What are temporal and spatial patterns of determines its functioning in some set of selection on integrated phenotypes in natural environmental conditions. The phenotypic populations? Are patterns of selection on characteristics may range from gene expression integrated phenotypes different than for profiles and metabolic fluxes to acoustic other traits?

5 • What are the genetic bases of phenotypic lineages maintain species cohesion? Can we plasticity in integrated phenotypes? How do develop a species concept for differences in the pattern and magnitude of and that captures plasticity affect selection and evolutionary their evolutionary history? responses? Evolution of conflict and cooperation Microbial Systems Scope: Conflict is pervasive in the evolution of Scope: Evolutionary biologists have devoted , and it is the key element of many major relatively little attention to microbial and viral transitions in the organization of life. systems, even though microorganisms make up Evolutionary conflict occurs in interactions most of the . Both systems are between species, between individuals, or particularly amenable to between genetic elements, and is sometimes and to comparative genomics. overcome by cooperation. Understanding how Rationale: New technologies that allow better cooperation arises is important in explaining sampling of both culturable and unculturable major transitions from the organization of genes genomes have expanded the tree of life, in , prokaryotes to , and uncovering a third domain, and have shown us other major transitions. the immense diversity in the microbial world. Rationale: Conflict and cooperation are crucial Evolutionary experiments using microbes or drivers of interactions within and between viruses have enriched our understanding of species. Between-species interactions include genetic change in novel environments, of host- plant-fungus interactions, pollination, host- evolution, and of the evolution of pathogen interactions, and ant-plant, or ant- conflict and cooperation. Medicine, agriculture fungus mutualisms. Selection operates on each and industry are increasingly using evolutionary partner to maximize their gain, and minimize approaches to microbial and viral systems to their cost, which can lead to arms races and interpret data, track , improve resulting rapid evolution. bioremediation, manage resistance, and design Within species interactions include male- and organisms for specific purposes. female interactions, worker-queen interactions, Microbial and viral systems are amenable to in mother-fetus interactions and -cell vivo reconstruction of ancestral genes and their interactions in chimeric organisms. Since the smaller genomes make them good candidates for partners are members of the same species, evolutionary genomics. These systems also selection for gain in one partner can favor genes provide a new testing ground for theories that are detrimental when expressed in the other developed for animals and plants, in which role, leading to overall evolutionary compromises horizontal gene transfer may be much less in the organism. For example, genes for traits important. optimal in males make worse females. Frontier questions What stabilizes these interactions, what • Are standard evolutionary models adequate allows cooperation to evolve in the face of to explain and predict microbial evolution? selfish genetic interests, and how the signature of To what extent do results from experimental competition remains are questions that have only studies of microbes and viruses generalize? recently been made accessible by advances in • How do selection and co-evolution operate genomics, , experimental in structured microbial communities such as evolution, and molecular manipulation. biofilms? Frontier questions • What are the mechanisms and evolutionary • What kinds of genes are important in social consequences of horizontal gene flow? How interactions? does horizontal gene transfer influence • How prevalent is the role of imprinting in cooperation and competition in microbial genomic conflicts? populations? • Is evolutionary conflict a major explanation • What is the diversity of genome structure for rapidly evolving genes? If so, what are among microbes and viruses, and what is the the pleiotropic effects of rapid evolution? functional significance of this diversity? • Are there general rules for the suppression • How can we quantify microbial and viral of conflict during the evolution of diversity, define species, and classify cooperation? How are cooperating systems unculturable organisms? How do microbial stabilized against cheaters?

6 • What determines the evolution of virulence? on evolution of phenotypic attributes within What is the genetic architecture of these groups. virulence? Frontier questions • What is the role of evolution of conflict and • Do different types of characters evolve at cooperation in major evolutionary different rates and at different phylogenetic transitions such as the origins of time scales within ? Are there regular multicellularity, endosymbiosis, and patterns of divergence for different types of chromosomal organization? phenotypic characters when compared • What is the role of sexual conflict in across clades? adaptation and speciation? • How does the environmental niche of species evolve and to what extent is a Large-Scale Patterns of Diversity in Time species’ range determined by adaptation to and Space changing conditions? Does evolutionary Scope: Large-scale patterns of , like change account for the success of invasive the latitudinal species gradient and the diversity species, and can such change be predicted? of communities, have not received satisfactory • How does the spatial structure of evolutionary explanations. Understanding the populations, species and communities affect evolutionary basis for these patterns will require evolutionary and coevolutionary change? synthesis of within-population, How does the spatial structure of populations, microevolutionary studies with across-taxa, species, and communities evolve? macroevolutionary investigations. • How can paleontological data be used to test Rationale: The availability of well-corroborated neontological hypotheses of selection and phylogenies, data bases of species occurrences, divergence? How can neontological data and global environmental data now document the test similar paleontological hypotheses? ecological and evolutionary scale of diversity at • How are processes of within-species a large scale and in great detail, and the divergence related to diversification of development of new conceptual approaches and clades? computational methods permits analysis of these data in manners not previously possible. Applied Evolution Phylogenetic studies have become the standard Scope: Applied evolution marshals our means of investigating historical, comparative understanding of evolutionary principles to questions, and a plethora of analytical methods tackle significant environmental, have been developed. At the same time, epidemiological, and agricultural problems. increasing understanding of past environments Rationale: The scale on which are and biotas permits the synthesis of neontological changing the in the 21st century is and paleontological data in novel and mutually unprecedented. Global pressures by humans on illuminating ways. Similarly, the recent planetary resources not only cause development of coalescent and comparative and detectable shifts in species’ geographic phylogeographic methods for inference of ranges and phenologies, but also cause within-species population histories now allows measurable evolutionary change in natural and similar questions to be investigated at the level introduced populations. Emerging agricultural of among-population differentiation. As a result, pests and pathogens are moved almost it is now possible to investigate the relationship instantaneously around the globe. Applied between processes leading to differentiation evolution seeks ways in which evolutionary among populations of the same species and those principles can improve the condition. Its leading to diversification of entire clades. applications range from improving ecosystem The challenge will be to develop methods services to developing better vaccines. It also that integrate across different disciplines and encompasses a variety of approaches for time-scales, spanning ecological, determining the impact of human activities on microevolutionary and macroevolutionary time. rates and patterns of evolutionary processes. The assembly of suitable large scale data sets is Finally, it includes efforts to trace the geographic vital. For example, the Tree-of-Life program is and taxonomic origins of rapidly spreading developing large-scale, highly resolved pathogens, to better understand their biological phylogenies for many groups of taxa; effects and predict further spread, and to devise comparable efforts are needed to collect the data means of control.

7 Examples of applied evolution span the • What is the impact of the agricultural entire breadth of evolutionary biology. Using the landscape on evolution of natural tools of DNA sequencing and phylogenetics, populations? evolutionary biologists can determine the time • What factors influence the extent and and geographic location of origin of invasive consequences of gene flow between GM species and emerging diseases. Evolutionary crops and natural populations? principles can be used to develop , ribozymes, and even organisms with specific functions and catalytic properties. Population INSTITUTIONAL genetics and ecological modeling can help RESOURCES predict if and how readily genes in genetically Many of the resources necessary to advance the modified organisms will spread into other frontiers of evolutionary biology are already in organisms. place. Others are only now coming into One major challenge in this field is to better existence. Some of the resources needed are predict the course of evolution given detailed institutional, but the scale of some questions will information on the genetics and behavior of require more collaboration among individual organisms. This aspect of applied evolution will investigators than has been typical of have its greatest successes in understanding evolutionary research in the past. responses of very simple systems, such as those • NESCent – The National Evolutionary in microbes or viruses. In many cases, these Synthesis Center will serve as a focal point same systems will be the ones with the greatest for a variety of activities including impact on human populations and environmental databasing and informatics development, change. catalysis of cross-disciplinary interactions, Basic research has much to offer here. and scientific synthesis. It will provide a Experimental evolutionary approaches, in which venue for the assembly, annotation, and evolutionary changes in laboratory populations integration of major evolutionary databases, are measured in real time, will play an important including genomic, phylogenetic, and role in understanding the diversity of genetic and phenotypic databases; for fostering the phenotypic responses to intense selection. By development of integrative, cross- providing biologists with a catalog of genes and disciplinary approaches to the solution of physiological properties, whole genome evolutionary questions; and for encouraging sequences of microbes and simple eukaryotes the integration and synthetic interpretation will also be a major bulwark of applied evolution of evolutionary data and ideas. in the future. Finally, large-scale monitoring of • NEON – The National Ecological environmental variables, such as those Observatory Network will provide an envisioned in NEON, will allow biologists to integrated, national platform at which a core predict with increasing certainty the spread and set of standardized ecological variables is fate of organisms and pathogens and their impact measured. It will provide a natural set of on agriculture and the environment. locations for large-scale experiments Frontier questions designed both to measure the strength, • Are there rules to predict changes in variability, and predictability of selection virulence or resistant of pathogens? Can we and to assess the influence of evolutionary predict potential pest/pathogen status based history on large-scale and on ecological, genomic or phylogenetic ecosystem processes. characteristics? • RCN – Research Collaboration Networks, • Can we use evolutionary theory to improve which had just been established at the time the efficacy of directed evolution? of the 1998 Working Group report, allow Conversely, can directed/experimental groups of individual investigators evolution inform evolutionary theory? investigating similar questions to coordinate • Can we predict changes in selection and their research. Investigators who take drift from changes in environmental advantage of the RCN program will be able conditions? to address more comprehensive questions • Can we predict the relative frequencies of than they could working in isolation from range shift, adaptation or in one another. response to environment change?

8 • FIBR – The Frontiers in Integrative populations should be a priority, including Biological Research program encourages non-model species that are close relatives of investigators to study “major, under-studied genetic model systems. The resources or unanswered questions in biology and needed are not only whole-genome to use innovative approaches to address sequences, but also the genomics tools that them by integrating the scientific concepts make genetic manipulation possible. To and research tools from across disciplines make these resources available, however, including biology, math and the physical evolutionary biologists will need to coalesce sciences, engineering, social sciences and around a selected set of evolutionary model the information sciences.” Many of the systems for genomic development. research frontiers identified in this report are • Phenomic resources: Evolutionary natural candidates for support from the analyses of complex, integrated phenotypes FIBR program. are limited by the capacity to measure and • IGERT – The Integrative Graduate analyze relevant phenotypes on large Education and Research Traineeship numbers of individuals in the context of program “is intended to catalyze a cultural evolutionary studies in the lab and field. change in graduate education … by Engineering resources for development of establishing innovative new models for new technologies for automated, high- graduate education and training.” Many of throughput phenotype measurement systems the research frontiers in this report require will be important to accomplish this task. integration of traditional biological • Large-scale experimental evolution: disciplines with the development of new Replicated, experimental studies of computational, statistical, and analytical evolution at a regional to national scale tools. Graduate training programs that seek would be invaluable for understanding to train such students are natural candidates short-term (1-100 year) evolutionary for support from the IGERT program. responses to environmental change in natural populations. The developing National Ecological Observatory Network INFRASTRUCTURAL NEEDS (NEON) holds great potential for enabling AND OPPORTUNITIES such large-scale evolutionary experiments. Much of the infrastructure required for Planning for instrumentation relevant to advancing evolutionary research is already in genetic and evolutionary analyses in the place, as outlined above. There are, however, context of NEON is needed to achieve this several areas in which the infrastructure falls potential. short and which are vital areas for future • Analytic resources: Databases for investment. phylogenetic, environmental, DNA • Genomic resources: The choice and sequence, and structure data are development of genomic data and tools to widely used and accessible, but comparable date have naturally focused on genetic resources for phenotypic data are lacking. model systems and species of applied The conceptual tools needed to develop an biomedical or agricultural importance. appropriate standard for phenotypic However, species vary in their utility for metadata have not been developed. Just as informing us about evolutionary patterns importantly, the computational and and processes. Mus musculus, Drosophila analytical tools necessary to integrate data melanogaster, Arabidopsis thaliana and derived from phylogenetic, environmental, other species for which extensive genomic DNA sequence, protein, and phenotypic data resources are now available allow us to have not been developed. Even tools for answer some evolutionary questions and visualization and analysis of variation in their full value is yet to be exploited, but complex phenotypes are largely lacking. these species are too divergent from one Advances in all of these areas are needed to another and represent too little evolutionary address many of the issues identified in this diversity to allow us to answer many critical report. questions in evolutionary biology. The development of genomic resources for evolutionary significant species and natural

9 Appendix: Workshop participants

Dr. Scott V. Edwards Dr. Therese Ann Markow Museum of Comparative Center for Insect Science Harvard University University of Arizona Cambridge, MA 02138 Tucson, AZ 85721 [email protected] [email protected]

Dr. Jack Hayes Dr. Patrick C. Phillips Biology Department Department of Biology University of Nevada Reno University of Oregon Reno, NV 89557 Eugene, OR 97403-5289 USA [email protected] [email protected]

Dr. Kent E. Holsinger Dr. Michael Purugganan Department of & Evolutionary Biology Department of Genetics University of Connecticut, U-3043 North Carolina State University Storrs, CT 06269-3043 Raleigh NC 27695 kent@.eeb.uconn.edu [email protected]

Dr. Scott Hodges Dr. Kerry L. Shaw Department of Ecology, Evolution & Marine Department of Biology Biology University of Maryland University of California College Park MD 20742-4415 Santa Barbara, CA 93106 [email protected] [email protected] Dr. Joan E. Strassmann Dr. Joel G. Kingsolver Dept. of Ecology and Evolutionary Biology, MS Department of Biology, CB-3280 170 University of North Carolina Rice University, 6100 Main St., Chapel Hill NC 27599-3280 Houston TX 77005-1892 USA [email protected] [email protected]

Dr. Jonathan B. Losos Dr. Holly A.Wichman Department of Biology Department of Biological Sciences Campus Box 1137 University of Idaho Washington University Moscow ID 83844-3051 Saint Louis, MO 63130-4899 [email protected] [email protected]

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