Annual Reviews OnlineSearch Annual Reviews OnlineAnnual Review ofEcology, Evolution,and Systematics OnlineMost Downloaded Annual Review of Ecology, Evolution,and Systematics ReviewsMost Cited Annual Review of Ecology, Evolution,and Systematics ReviewsAnnual Review of Ecology, Evolution,and Systematics ErrataView Current Editorial CommitteeAll Articles in the Annual Review of Ecology, Evolution, and Systematics, Vol. 39Top Predators as Conservation Tools: Ecological Rationale, Assumptions, and EfficacyRevisiting the Impact of Inversions in Evolution: From Population Genetic Markers to Drivers of Adaptive Shifts and Speciation?Radial Symmetry, the Anterior/Posterior Axis, and Echinoderm Hox GenesThe Great American Schism: Divergence of Marine Organisms After the Rise of the Central American IsthmusThe Ecological Performance of Protected AreasMorphological Integration and Developmental ModularityHerbivory from Individuals to EcosystemsStoichiometry and Nutrition of Plant Growth in Natural CommunitiesPlague Minnow or Mosquito Fish? A Review of the Biology and Impacts of Introduced Gambusia SpeciesThe Impact of Natural Selection on the Genome: Emerging Patternsin Drosophila and ArabidopsisSanctions, Cooperation, and the Stability of Plant-Rhizosphere MutualismsShade Tolerance, a Key Plant Feature of Complex Nature and ConsequencesThe Impacts of Fisheries on Marine Ecosystems and the Transition to Ecosystem-Based ManagementThe Performance of the Endangered Species ActPhylogenetic Approaches to the Study of ExtinctionPhylogenetic Approaches to the Study of ExtinctionAdaptation to Marginal HabitatsConspecific Brood Parasitism in Birds: A Life-History PerspectiveStratocladistics: Integrating Temporal Data and Character Data in Phylogenetic InferenceThe Evolution of Animal WeaponsUnpacking β: Within-Host Dynamics and the Evolutionary Ecology of Pathogen TransmissionEvolutionary Ecology of Figs and Their Associates: Recent Progress and Outstanding PuzzlesThe Earliest Land PlantsSpatial Dynamics of FoodwebsSpecies Selection: Theory and DataNew Answers for Old Questions: The Evolutionary Quantitative Genetics of Wild Animal PopulationsWake Up and Smell the Roses: The Ecology and Evolution of Floral ScentEver Since Owen: Changing Perspectives on the Early Evolution of TetrapodsPandora’s Box Contained Bait: The Global Problem of Introduced EarthwormsTrait-Based Community Ecology of PhytoplanktonWhat Limits Trees in C4 Grasslands and Savannas?ANRV360-ES39-21 ARI 1 October 2008 13:30Evolutionary Ecology of Figsand Their Associates:Recent Progress andOutstanding PuzzlesEdward Allen Herre,1K. Charlotte Jand´er,1,2and Carlos Alberto Machado31Smithsonian Tropical Research Institute, Apartado 0843-03092, Balboa, Ancon, Republic ofPanama; email: [email protected] of Neurobiology and Behavior, Cornell University, Ithaca, New York 148533Department of Biology, University of Maryland, College Park, Maryland 20742Annu. Rev. Ecol. Evol. Syst. 2008. 39:439–58First published online as a Review in Advance onSeptember 2, 2008The Annual Review of Ecology, Evolution, andSystematics is online at ecolsys.annualreviews.orgThis article’s doi:10.1146/annurev.ecolsys.37.091305.110232Copyrightc 2008 by Annual Reviews.All rights reserved1543-592X/08/1201-0439$20.00Key Wordsbreeding systems, coevolution, frugivory, mutualism, parasitism,pollination, sex allocation, tradeoffsAbstractOver the past decade a proliferation of research has enriched and dramati-cally altered our understanding of the biology of figs, their pollinator wasps,and the myriad of other organisms that depend on them. Ecologically, thiswork underscores the crucial role that fig fruits play in sustaining and shapingtropical frugivore communities. More generally, this work addresses severalkey issues in evolutionary ecology, including evolution of breeding systems(shifts between monoecy and dioecy), factors that promote the stability ofmutualisms, precision of adaptation, and trajectories of community assem-bly and coevolution in systems with multiple interacting partners. Moreover,both the pollinating and nonpollinating wasps associated with figs provideunparalleled opportunities for examining how different population struc-tures can differentially affect sex allocation, kin selection, the evolution ofparasite virulence, and many fundamental parameters of population genet-ics (e.g., levels of genetic variation and rates of silent and nonsilent basesubstitutions).439Annu. Rev. Ecol. Evol. Syst. 2008.39:439-458. Downloaded from arjournals.annualreviews.orgby UNIVERSITY OF MARYLAND - COLLEGE PARK - MCKELDIN LIBRARY on 03/03/10. For personal use only.Click here for quick links to Annual Reviews content online, including:• Other articles in this volume• Top cited articles• Top downloaded articles• Our comprehensive searchFurtherANNUALREVIEWSANRV360-ES39-21 ARI 1 October 2008 13:30“If you study tropical frugivores, or carnivores for that matter, directly or indirectly you will end up studying figs.”-Charles Handley, Curator of Birds and Mammals, National Museum of Natural History, Washington,DC, 1950–2000“If you study figs, you will end up studying wasps.” -J.T. Wiebes, Curator of the Museum of Natural Historyand Chairman of the Department of Zoology, Leiden, Netherlands, 1954–1998“If you study the wasps, you will end up studying their figs.” -Finn Kjellberg, current Directeur de Rechercheau Centre d’Ecologie Fonctionnelle et Evolutive, Montpellier, France“If you study figs, you will end up studying the animals that eat them” -VariousFigs (Ficus spp.) depend on wasps for transmission of their pollen and, therefore, viable seedproduction, whereas fig wasps depend on fig inflorescences for the completion of their life cycle.This relationship, fundamental to the reproduction of both parties, has extraordinary ecologicalconsequences. From pea- to peach-sized or larger, the fruits of different fig species may ripenred, yellow, green, or purple, and may attract any of a wide range of frugivores, by either visualor chemical signaling (Berg & Wiebes 1992, Borges et al. 2008, Corner 1952, Harrison 2005,Kalko et al. 1996). Figs are truly keystone resources that provide the metabolic foundation forthe pollinator and parasitic wasps, nematodes, mites, etc., that live within the microcosm of theirfruit (Corner 1952, Compton et al. 1994). Figs also provide keystone resources at larger scales forcommunities of