Evolutionary Dynamics of Seed Size and Seedling Competitive Ability Geritz, S.A.H., Meijden, E. van der and Metz, J.A.J. IIASA Interim Report October 1997 Geritz, S.A.H., Meijden, E. van der and Metz, J.A.J. (1997) Evolutionary Dynamics of Seed Size and Seedling Competitive Ability. IIASA Interim Report. IIASA, Laxenburg, Austria, IR-97-071 Copyright © 1997 by the author(s). http://pure.iiasa.ac.at/5224/ Interim Reports on work of the International Institute for Applied Systems Analysis receive only limited review. Views or opinions expressed herein do not necessarily represent those of the Institute, its National Member Organizations, or other organizations supporting the work. All rights reserved. Permission to make digital or hard copies of all or part of this work for personal or classroom use is granted without fee provided that copies are not made or distributed for profit or commercial advantage. All copies must bear this notice and the full citation on the first page. For other purposes, to republish, to post on servers or to redistribute to lists, permission must be sought by contacting [email protected] IIASA International Institute for Applied Systems Analysis • A-2361 Laxenburg • Austria Tel: +43 2236 807 • Fax: +43 2236 71313 • E-mail: [email protected] • Web: www.iiasa.ac.at INTERIM REPORT IR-97-071/October Evolutionary Dynamics of Seed Size and Seedling Competitive Ability S.A.H. Geritz ([email protected]) E. van der Meijden ([email protected]) J.A.J. Metz ([email protected]) Approved by Ulf Dieckmann ([email protected]) Project Coordinator, Adaptive Dynamics Network Interim Reports on work of the International Institute for Applied Systems Analysis receive only limited review. Views or opinions expressed herein do not necessarily represent those of the Institute, its National Member Organizations, or other organizations supporting the work. IIASA STUDIES IN ADAPTIVE DYNAMICS ADN NO. 19 The Adaptive Dynamics Network at IIASA fosters the development of new mathematical and conceptual techniques for understanding the evolution of complex adaptive systems. Focusing on these long-term implications of adaptive processes in systems of limited growth, the Adaptive Dynamics Network brings together scientists and institutions from around the world with IIASA acting as the central node. Scientific progress within the network is reported in the IIASA Studies in Adaptive Dynamics series. THE ADAPTIVE DYNAMICS NETWORK The pivotal role of evolutionary theory in life sciences derives from its capability to provide causal explanations for phenomena that are highly improbable in the physicochemical sense. Yet, until recently, many facts in biology could not be accounted for in the light of evolution. Just as physicists for a long time ignored the presence of chaos, these phenomena were basically not perceived by biologists. Two examples illustrate this assertion. Although Darwin’s publication of “The Origin of Species” sparked off the whole evolutionary revolution, oddly enough, the population genetic framework underlying the modern synthesis holds no clues to speciation events. A second illustration is the more recently appreciated issue of jump increases in biological complexity that result from the aggregation of individuals into mutualistic wholes. These and many more problems possess a common source: the interactions of individuals are bound to change the environments these individuals live in. By closing the feedback loop in the evolutionary explanation, a new mathematical theory of the evolution of complex adaptive systems arises. It is this general theoretical option that lies at the core of the emerging field of adaptive dynamics. In consequence a major promise of adaptive dynamics studies is to elucidate the long-term effects of the interactions between ecological and evolutionary processes. A commitment to interfacing the theory with empirical applications is necessary both for validation and for management problems. For example, empirical evidence indicates that to control pests and diseases or to achieve sustainable harvesting of renewable resources evolutionary deliberation is already crucial on the time scale of two decades. The Adaptive Dynamics Network has as its primary objective the development of mathematical tools for the analysis of adaptive systems inside and outside the biological realm. IIASA STUDIES IN ADAPTIVE DYNAMICS No. 1 Metz JAJ, Geritz SAH, Meszéna G, Jacobs FJA, van Heerwaarden JS: Adaptive Dynamics: A Geometrical Study of the Consequences of Nearly Faithful Reproduction. IIASA Working Paper WP-95-99. van Strien SJ, Verduyn Lunel SM (eds.): Stochastic and Spatial Structures of Dynamical Systems, KNAW Verhandelingen, North Holland, Amsterdam, pp. 183-231 (1996). No. 2 Dieckmann U, Law R: The Dynamical Theory of Coevolution: A Derivation from Stochastic Ecological Processes. IIASA Working Paper WP-96-01. Journal of Mathematical Biology (1996) 34, 579–612. No. 3 Dieckmann U, Marrow P, Law R: Evolutionary Cycling of Predator-Prey Interactions: Population Dynamics and the Red Queen. Journal of Theoretical Biology (1995) 176, 91–102. No. 4 Marrow P, Dieckmann U, Law R: Evolutionary Dynamics of Predator-Prey Systems: An Ecological Perspective. IIASA Working Paper WP-96-02. Journal of Mathematical Biology (1996) 34, 556–578. No. 5 Law R, Marrow P, Dieckmann U: On Evolution under Asymmetric Competition. IIASA Working Paper WP-96-03. Evolutionary Ecology (1997) 11, 485–501. No. 6 Metz JAJ, Mylius SD, Diekmann O: When Does Evolution Optimise? On the Relation between Types of Density Dependence and Evolutionarily Stable Life History Parameters. IIASA Working Paper WP-96-04. No. 7 Ferrière R, Gatto M: Lyapunov Exponents and the Mathematics of Invasion in Oscillatory or Chaotic Populations. Theoretical Population Biology (1995) 48, 126–171. No. 8 Ferrière R, Fox GA: Chaos and Evolution. Trends in Ecology and Evolution (1995) 10, 480–485. No. 9 Ferrière R, Michod RE: The Evolution of Cooperation in Spatially Heterogeneous Populations. IIASA Working Paper WP-96-29. American Naturalist (1996) 147, 692–717. No. 10 Van Dooren TJM, Metz JAJ: Delayed Maturation in Temporally Structured Populations with NonEquilibrium Dynamics. IIASA Working Paper WP-96-70. Journal of Evolutionary Biology (1997) in press. No. 11 Geritz SAH, Metz JAJ, Kisdi E, Meszéna G: The Dynamics of Adaptation and Evolutionary Branching. IIASA Working Paper WP-96-77. Physical Review Letters (1997) 78, 2024–2027. No. 12 Geritz SAH, Kisdi E, Meszéna G, Metz JAJ: Evolutionarily Singular Strategies and the Adaptive Growth and Branching of the Evolutionary Tree. IIASA Working Paper WP-96-114. Evolutionary Ecology (1997) in press. No. 13 Heino M, Metz JAJ, Kaitala V: Evolution of Mixed Maturation Strategies in Semelparous Life-Histories: the Crucial Role of Dimensionality of Feedback Environment. IIASA Working Paper WP-96-126. Philosophical Transactions of the Royal Society of London Series B (1997) in press. No. 14 Dieckmann U: Can Adaptive Dynamics Invade? IIASA Working Paper WP-96-152. Trends in Ecology and Evolution (1997) 12, 128–131. No. 15 Meszéna G, Czibula I, Geritz SAH: Adaptive Dynamics in a Two-Patch Environment: a Simple Model for Allopatric and Parapatric Speciation. IIASA Interim Report IR-97-01. Journal of Biological Systems (1997) in press. No. 16 Heino M, Metz JAJ, Kaitala V: The Enigma of Frequency-Dependent Selection. IIASA Interim Report IR-97-61. No. 17 Heino M: Management of Evolving Fish Stocks. IIASA Interim Report IR-97-62. No. 18 Heino M: Evolution of Mixed Reproductive Strategies in Simple Life-History Models. IIASA Interim Report IR-97-63. No. 19 Geritz SAH, van der Meijden E, Metz JAJ: Evolutionary Dynamics of Seed Size and Seedling Competitive Ability. IIASA Interim Report IR-97-71. No. 20 Galis F, Metz JAJ: Why are there so many Cichlid Species? On the Interplay of Speciation and Adaptive Radiation. IIASA Interim Report IR-97-72. Trends in Ecology and Evolution (1997) in press. No. 21 Boerlijst MC, Nowak MA, Sigmund K: Equal Pay for all Prisoners. / The Logic of Contrition. IIASA Interim Report IR-97-73. AMS Monthly (1997) 104, 303–307. Theoretical Biology (1997) 185, 281–294. No. 22 Law R, Dieckmann U: Symbiosis without Mutualism and the Merger of Lineages in Evolution. IIASA Interim Report IR-97-74. No. 23 Klinkhamer PGL, de Jong TJ, Metz JAJ: Sex and Size in Cosexual Plants. IIASA Interim Report IR-97-78. Trends in Ecology and Evolution (1997) 12, 260–265. Issues of the IIASA Studies in Adaptive Dynamics series can be obtained free of charge. Please contact: Adaptive Dynamics Network International Institute for Applied Systems Analysis Schloßplatz 1 A–2361 Laxenburg Austria Telephone +43 2236 807, Telefax +43 2236 71313, E-Mail [email protected], Internet http://www.iiasa.ac.at/Research/ADN Contents ,QWURGXFWLRQ (YROXWLRQDULO\6LQJXODU6WUDWHJLHV 5HVLGHQWSRSXODWLRQG\QDPLFVDQGPXWDQW¶VILWQHVV (YROXWLRQDULO\VWDEOHVLQJXODUVWUDWHJLHV &RQYHUJHQFHVWDEOHVLQJXODUVWUDWHJLHV (YROXWLRQDU\EUDQFKLQJ &KDQJHVLQQXPEHUDQGVWDELOLW\SURSHUWLHVRIHYROXWLRQDU\VLQJXODUVWUDWHJLHV (YROXWLRQDULO\6LQJXODU&RDOLWLRQV 5HVLGHQWSRSXODWLRQG\QDPLFVDQGPXWDQW¶VILWQHVV ,VRFOLQHVDQGHYROXWLRQDULO\VLQJXODUFRDOLWLRQV &KDQJHLQQXPEHUDQGVWDELOLW\SURSHUWLHVRIHYROXWLRQDU\VLQJXODUFRDOLWLRQV *HQHUDOL]DWLRQWRKLJKHUOHYHOVRISRO\PRUSKLVP ([WLQFWLRQRIHYROXWLRQDU\EUDQFKHV &RQFOXVLRQVDQG'LVFXVVLRQ $SSHQGL[ ,VRFOLQHVDQGWKHLUFRQQHFWLRQWRWKHERXQGDU\RIWKHDUHDRIFRH[LVWHQFH /RVVRIJOREDOHYROXWLRQDU\VWDELOLW\DQGWKHHPHUJHQFHRIQHZVLQJXODULWLHV 5HIHUHQFHV Abstract :H SUHVHQW D PRGHO IRU WKH HYROXWLRQDU\ G\QDPLFV RI VHHG VL]H ZKHQ VHHGOLQJV IURP ODUJHVHHGVDUHEHWWHUFRPSHWLWRUVWKDQVHHGOLQJVIURPVPDOOVHHGVDQGWKHUHLVDWUDGH RIIEHWZHHQVHHGVL]HDQGVHHGQXPEHU:HILUVWFRQVLGHUWZROLPLWLQJFDVHVZKHUHVHHG VL]HKDVHLWKHUQRHIIHFWRQWKHFRPSHWLWLYHDELOLW\RIVHHGOLQJVRUZKHUHVHHGOLQJVIURP ODUJHU VHHGV DOZD\V ZLQ IURP VHHGOLQJV IURP VPDOOHU VHHGV LI WRJHWKHU LQ WKH VDPH JHUPLQDWLRQ VLWH ,Q WKH ILUVW FDVH WKHUH LV D VLQJOH HYROXWLRQDU\ RSWLPDO VHHG VL]H H[FOXGLQJ DOO RWKHU ZKHUHDV LQ WKH VHFRQG FDVH WKHUH LV DQ HYROXWLRQDULO\ VWDEOH VHHG SRO\PRUSKLVP ZLWK D FRQWLQXRXV YDULDWLRQ RI VHHG VL]HV ZKHUH SODQWV ZLWK VPDOO EXW QXPHURXVVHHGVVXUYLYHE\H[SORLWLQJVLWHVWKDWE\FKDQFHUHPDLQXQRFFXSLHGE\SODQWV ZLWKODUJHUEXWOHVVQXPHURXVVHHGV:HLQYHVWLJDWHKRZ WKHVH WZR FDVHV FRQQHFW WR RQH DQRWKHU YLD LQWHUPHGLDWH OHYHOV RI FRPSHWLWLYH DV\PPHWU\ :H ILQG WKDW VWURQJ FRPSHWLWLYH DV\PPHWU\ DQG KLJK UHVRXUFH OHYHOV IDYRU FRH[LVWHQFH RI SODQWV ZLWK GLIIHUHQWVHHGVL]HVZKHQVHHGDQGVHHGOLQJVXUYLYDOLVPRGHUDWHO\ORZEXWODUJHVHHGV KDYH D VXEVWDQWLDO SUHFRPSHWLWLYH DGYDQWDJH RYHU VPDOOHU VHHGV $VVXPLQJ PXWDWLRQ OLPLWHG HYROXWLRQ DQG DVVXPLQJ WKDW VLQJOH PXWDWLRQV KDYH RQO\ D VPDOO SKHQRW\SLF HIIHFWDQLQLWLDOO\PRQRPRUSKLFSRSXODWLRQZLWKDVLQJOHVHHGVL]HZLOOUHDFKWKHILQDO HYROXWLRQDULO\ VWDEOH SRO\PRUSKLF VWDWH WKURXJK D VHULHV RI GLVFUHWH HYROXWLRQDU\ EUDQFKLQJ HYHQWV $W HDFK EUDQFKLQJ HYHQW D JLYHQ OLQHDJH DOUHDG\ SUHVHQW LQ WKH SRSXODWLRQ GLYLGHV LQWR WZR GDXJKWHU OLQHV HDFK ZLWK LWV RZQ VHHG VL]H ,I SUHFRPSHWLWLYH VHHG DQG VHHGOLQJ VXUYLYDO LV KLJK IRU VPDOO DQG ODUJH VHHGV DOLNH HYROXWLRQDU\ EUDQFKLQJ PD\ EH IROORZHG E\ H[WLQFWLRQ RI RQH RU PRUH OLQHDJHV LQFOXGLQJPDVVH[WLQFWLRQDQGWKXVQRWQHFHVVDULO\JLYHVULVHWRHYROXWLRQDULO\VWDEOH VHHG SRO\PRUSKLVPV 9DULRXV UHVXOWV SUHVHQWHG KHUH DUH PRGHOLQGHSHQGHQW DQG SRLQW WKH ZD\ WR D PRUH JHQHUDO HYROXWLRQDU\ ELIXUFDWLRQ WKHRU\ GHVFULELQJ KRZ WKH QXPEHU DQG VWDELOLW\ SURSHUWLHV RI HYROXWLRQDU\ HTXLOLEULD FDQ FKDQJH DV D FRQVHTXHQFH RI FKDQJHVLQPRGHOSDUDPHWHUV .H\ZRUGV 6HHG VL]H HYROXWLRQ VHHGOLQJ FRPSHWLWLRQ IUHTXHQF\GHSHQGHQW VHOHFWLRQ DGDSWLYHG\QDPLFVHYROXWLRQDU\EUDQFKLQJH[WLQFWLRQHYROXWLRQDU\ELIXUFDWLRQWKHRU\ About the Authors 6$+*HULW] 'HSDUWPHQWRI=RRORJ\ 8QLYHUVLW\RI0DU\ODQG &ROOHJH3DUN0DU\ODQG 86$ DQG ,QVWLWXWHRI(YROXWLRQDU\DQG(FRORJLFDO6FLHQFHV .DLVHUVWUDDW *3/HLGHQ 7KH1HWKHUODQGV (YDQGHU0HLMGHQ ,QVWLWXWHRI(YROXWLRQDU\DQG(FRORJLFDO6FLHQFHV .DLVHUVWUDDW *3/HLGHQ 7KH1HWKHUODQGV -$-0HW] ,QVWLWXWHRI(YROXWLRQDU\DQG(FRORJLFDO6FLHQFHV .DLVHUVWUDDW *3/HLGHQ 7KH1HWKHUODQGV DQG $GDSWLYH'\QDPLFV1HWZRUN ,QWHUQDWLRQDO,QVWLWXWHIRU$SSOLHG6\VWHP$QDO\VLV_ /D[HQEXUJ$XVWULD Acknowledgments 7KLV ZRUN ZDV VXSSRUWHG E\ JUDQWV IURP WKH 1HWKHUODQGV 2UJDQL]DWLRQ IRU 6FLHQWLILF 5HVHDUFK1:2DQGIURPWKH8QLYHUVLW\RI0DU\ODQGDW&ROOHJH3DUN:HWKDQNeYD .LVGL3HWHU.OLQNKDPHU7RPGH-RQJ*p]D0HV]pQD)UDQV-DFREVDQG3HWHU$EUDPV IRUPDQ\GLVFXVVLRQVRQWKHWRSLFRIVHHGVL]HHYROXWLRQDQGHYROXWLRQDU\G\QDPLFV Evolutionary Dynamics of Seed Size and Seedling Competitive Ability S.A.H. 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