evolutionary demography society 2015...welcome!(! 1!...

Evolutionary Demography Society 2015

Program and Abstracts

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The$COMPADRE$Plant$Matrix$Database,$and$the$new$COMADRE$Animal$Matrix$Database,$hosted$at$the$Max$Planck$Institute$for$Demographic$Research,$make$demographic$data$available$for$many$species.$$New$developments$will$be$presented$by$Rob$SalgueroGGomez$GGG$a$talk$and$poster$Monday'afternoon;$a$workshop$on$Wednesday'afternoon.$

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Welcome!(

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Welcome!to!the!2015!meeting!of!the!Evolutionary!Demography!Society.!This!is!our!third!meeting,!following!the!2014!meeting!at!Stanford!University!and!the!inaugural!2013!meeting!at!the!University!of!Southern!Denmark.!The!2016!meeting!will!be!held!at!the!University!of!Virginia.!Our!meetings!alternate!between!Europe!and!North!America.!!We!hope!you!enjoy!the!conference.!The!diverse1!program!

contains!some!20Lminute!“keynote”!talks,!but!most!presentations!are!Lightning!Talks!of!5!minutes!(plus!5!minutes!for!questions).!It’s!a!surprisingly!good!way!to!communicate.!There!are!posters;!take!advantage!of!the!chance!to!talk!to!people!about!their!research.!The!Congrescentrum!de!Werelt!has!excellent!facilities!for!informal!discussions;!enjoy!them.!!!Join(the(Society:!Are!you!an!evolutionary!demographer?!If!you!are!reading!this,!the!chances!are!good!that!you!are.!So!you!should!join!the!Society.!Visit!our!web!site!at:!!

www.evodemos.org!!Thank(you(…(We!would!particularly!like!to!acknowledge!the!Institute!for!Biodiversity!and!Ecosystem!Dynamics,!University!of!Amsterdam,!for!financial!assistance!with!the!conference.!!!Organizing!committee:!Nienke!Hartemink!and!Hal!Caswell!!Cover!image:!Karel!Appel,!Mens%en%Dieren%(Man%and%Animals),!1949.!Permission:!Collection!Stedelijk!Museum,!Amsterdam.!!Karel!Appel!(1921L2006)!was!a!Dutch!painter,!sculptor,!and!poet.!His!painting!Mens%en%Dieren!is!on!display!in!the!Stedelijk!Museum!in!Amsterdam.!It!is!huge,!playful,!and!well!worth!a!visit.!And!it!perfectly!captures!the!spirit!of!the!Evolutionary!Demography!Society.!(Apologies!to!the!plant!demographers;!for!the!moment,!we!will!consider!plants!as!a!kind!of!animal.)!!

!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!1!The!Wordle!cloud!became!more!diverse!after!removing!the!overwhelmingly!dominant!word!“population.”!!

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Monday,(5(October(2015(

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Monday!5th!of!October!2015!! Presenter!! Title!(abridged)!8:50!–10:00! Registration,(coffee(and(tea(

10:00L10:20! Welcome!speech! ! !10:20L10:50! Keynote%1:!Hal!Caswell! How!heterogeneity!works:!an!outline!of!a!

general!demographic!theory!10:50L11:00! Simon!Verhulst! A!new!tool,!s,!to!quantify!the!contribution...!11:00L11:10! Silke!van!Daalen! Variation!in!lifetime!reproduction!...!11:10L11:20! Célia!Rezouki! SociallyLmediated!effects!of!climate!...!11:20L11:30! Lynn!Govaert! EcoLevolutionary!partitioning!metrics!...!11:30L12:00! Coffee(break(

12:00L12:10! Hanna!ten!Brink! Evolution!of!metamorphosis...!12:10L12:20! Mollie!Brooks! StateLspace!autoregressive!growth...!!12:20L12:30( Paul!van!de!Vijver! Early!and!extraordinary!physical!peaks!...!12:30L13:30! Lunch(

13:30L14:00! Keynote%2:!James!Holland!Jones! TBA!!

14:00L14:10! Ulrich!Steiner! Sensitivity!of!the!entropy!...!!14:10L14:20! Monique!de!Jager! LabyrinthLlike!population!structures...!14:20L14:30! Andrea!Westerband! Physiological!responses!to!light...!14:30L14:40! Owen!Jones! Exploring!the!evolution!of!life!history!...!14:40L14:50! Masahito!Morita! Fertility!decline!in!Japan!...!14:50L15:00! David!Sharrow! Left!truncation!in!human!survival!modeling!

15:00L15:30!Keynote%3:!Andre!de!Roos! Nonlinear!evolutionary!demography!of!

complex!life!cycles!15:30L16:00! Coffee(break(

16:00L16:10! Morgane!Tidière! Variability!of!actuarial!senescence!...!16:10L16:20! Chloe!Nater! Exogenous!and!endogenous!...!!16:20L16:30! Luke!EberhartLPhillips! Consequences!of!stageL!and!sexLspecific!...!16:30L16:40! Beate!Proske! EcoLevolutionary!dynamics!...!!16:40L16:50! Charlotte!de!Vries! Demography!when!history!matters!...!16:50L17:10! Short(break(

17:10L17:20! Anne!Kwak! Does!sampling!methodology!affect!...!!17:20L17:30! Diana!Rypkema! Impacts!of!interaction!between!...!17:30L17:40! Tom!Miller! Effects!of!microbial!symbionts!...!17:40L17:50! Rob!SalgueroLGomez! The!COMADRE!animal!matrix!database!...!17:50L18:20! Drinks(+(posters(

18:30! Dinner(

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! 3!

How(heterogeneity(works:(an(outline(of(a(general(demographic(theory!!

Hal!Caswell!Institute!for!Biodiversity!and!Ecosystem!Dynamics!University!of!Amsterdam!

Accounting!for!heterogeneity!(differences!among!individuals!in!their!vital!rates)!is!the!oldest!problem!in!demography.!When!the!heterogeneity!is!not!accounted!for,!it!can!distort!the!conclusions!based!on!demographic!models.!!Heterogeneity!is!tamed!by!accounting!for!it;!doing!so!reduces!problem!to!that!of!stageLclassified!demography.!!We!now!possess!most!of!the!pieces!needed!for!a!general!theory!of!heterogeneity,!based!on!the!vecLpermutation!matrix!formulation!of!multistate!models.!!The!theory!relies!on!four!sets!of!matrices,!defining!fertility,!the!dynamics!of!demographic!stages!and!heterogeneity!groups,!and!the!transmission!of!heterogeneity.!!It!permits!calculation!of,!inter%alia,!the!moments!of!longevity,!measures!of!discrepancy,!the!distribution!of!age!or!stage!at!death,!quasiLstationary!distributions,!population!growth!rates!and!structure,!and!the!sensitivity!and!elasticity!of!all!these!quantities!to!changes!in!the!vital!rates!and!the!parameters!defining!the!transmission!of!heterogeneity.!!It!operates!equally!at!the!cohort!and!the!population!levels,!and!makes!it!possible!to!calculate!variance!due!to!individual!stochasticity,!and!population!growth!effects!due!to!demographic!stochasticity.!!Fortunately,!however,!not!all!the!problems!relating!to!heterogeneity!have!been!solved.!!!A(new(tool,(s,(to(quantify(the(contribution(of(senescence(to(lifespan((Simon!Verhulst1,!Ferdinand!Verhulst2!&!Rampal!Etienne1!

!1Groningen%Institute%of%Evolutionary%Life%Sciences,%University%of%Groningen,%the%Netherlands%

2%Mathematical%Institute,%Utrecht%University,%the%Netherlands%

!Different!combinations!of!the!Gompertz!equation!parameters!α!and!β!(in!mortality!rate!μ(x)=αeβx,!where!x!is!age)!can!yield!the!same!life!span.!In!this!equation,!α!determines!the!vulnerability!to!senescence,!while!β!determines!the!rate!of!senescence.!In!the!unusual!case!that!the!average!lifespan!of!two!populations!is!identical,!the!contribution!of!actuarial!senescence!to!determining!average!lifespan!can!be!compared!using!β.!However,!in!the!more!usual!case,!where!average!lifespan!of!two!populations!is!not!identical,!the!contribution!of!senescence!to!lifespan!is!more!difficult!to!compare.!To!this!end!we!propose!a!new!metric,!s,!which!is!the!slope!of!the!relation!between!α!and!β!in!isoLlifespan!plots.!We!use!s!to!investigate!how!the!contribution!of!senescence!to!lifespan!varies!between!populations!of!animals!and!humans.!Preliminary!results!from!animal!and!human!studies!suggest!increased!lifespan!is!achieved!primarily!through!lower!vulnerability!to!senescence.!!Variation(in(lifetime(reproduction:(individual(stochasticity(and(heterogeneity(

Silke!van!Daalen!and!Hal!Caswell!!Institute%for%Biodiversity%and%Ecosystem%Dynamics%(IBED),%University%of%Amsterdam,%the%Netherlands%

!Variation!in!lifetime!reproduction!in!many!species!is!often!interpreted!as!a!measure!of!potential!for!natural!selection.!However,!this!interpretation!is!only!valid!if!the!variation!in!question!is!heritable.!It!is!possible!to!separate!underlying!sources!of!variation!into!heterogeneity!and!individual!stochasticity,!where!heterogeneity!refers!to!genuine!individual!differences,!and!individual!stochasticity!refers!to!the!randomness!in!the!survival!and!reproduction!of!individuals!experiencing!identical!rates.!In!our!analyses,!we!have!calculated!the!standardized!variance,!representing!the!opportunity!for!selection,!accounted!for!by!individual!stochasticity,!and!we!compare!that!with!the!empirical!measured!standardized!variance.!We!use!ageL!and!stageLclassified!demographic!data!for!a!range!of!mammal!and!bird!species!and!analytically!calculate!the!variation!in!lifetime!reproductive!output!using!Markov!chains!with!rewards.!We!show!for!a!range!of!species!that!most,!and!in!some!cases!all!of!the!variation!in!lifetime!reproduction!is!due!to!individual!stochasticity.!In!order!to!link!variation!to!an!opportunity!for!selection,!one!must!calculate!the!contribution!of!individual!stochasticity!to!the!variation!in!lifetime!reproductive!output.!!


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SociallyOmediated(effects(of(climate(change(decrease(survival(of(the(hibernating(Alpine(marmots((

Célia!Rézouki1!,!Marion!Tafani1,!Aurélie!Cohas1,!Anne!Loison2,!JeanLMichel!Gaillard1,!Dominique!Allainé1!and!Christophe!Bonenfant1!!

1%Université%de%Lyon,%FH69000,%Lyon;%Université%Lyon%1;%CNRS%UMR5558,%Laboratoire%de%Biométrie%et%Biologie%

Évolutive,%FH69622,%Villeurbanne,%France.%2%Laboratoire%d’Écologie%Alpine,%CNRS%UMR5553,%Université%de%Savoie,%Bâtiment%Belledonne,%FH73376,%Le%

BourgetHduHLac,%France.%

!! An!increasing!challenge!in!ecological!research!is!to!explain!betweenLspecies!differences!in!their!response!to!climate!change,!which!includes!understanding!the!way!climate!shapes!species!life!history!variations.!We!propose!that!speciesLspecific!life!history!features,!particularly!those!that!are!associated!with!environmental!constraints,!such!as!hibernation!and!sociality,!should!condition!the!way!climate!influences!population!dynamics.!To!test!our!hypothesis!we!use!a!23!yearLlong!monitoring!on!an!Alpine!marmot!population!in!the!French!Alps,!to!investigate!how!this!species'!lifestyle,!more!specifically,!how!being!an!hibernator!and!displaying!a!high!degree!of!sociality,!interacts!with!the!ongoing!climate!change!in!shaping!ageLspecific!survival!pattern.!! We!find!clear!evidence!for!a!keyLrole!of!the!lifestyle!in!the!response!to!climate!change!of!the!Alpine!marmot.!We!demonstrate!that!juvenile!survival!strongly!decrease!over!time!because!of!interLrelated!effects!of!social!factors!and!winter!weather!conditions.!In!the!Alpine!marmot,!helpers!increase!juvenile!survival!by!buffering!them!against!the!energetic!cost!of!hibernation.!Over!time,!harsher!winters!lower!the!juvenile!survival,!which!in!turn!decreases!the!relative!number!of!helpers!present!in!the!population!in!the!following!years!and!therefore!the!chances!of!juveniles!born!in!later!years!to!survive!over!their!first!winter.!This!mechanism!we!uncover!indicates!that!the!Alpine!marmot!may!have!entered!a!vicious!circle!due!to!climate!change,!in!relation!with!the!constraints!associated!with!its!sociality.!! Our!findings!contrast!with!responses!to!climate!change!reported!to!date!for!hibernating!species,!like!the!yellowLbellied!marmot,!a!closelyLrelated!species!that!do!not!necessarily!hibernate!in!family!groups,!and!currently!benefits!from!the!ongoing!climate!change.!Such!contrasting!responses!suggest!that!the!species!lifestyle!might!be!involved!in!betweenLspecies!differences!in!lifeLhistory!responses!to!climate!change,!an!aspect!that!has!been!overlooked!so!far.!!

!EcoOevolutionary(partitioning(metrics:(assessing(the(importance(of(ecology(and(evolution(as(drivers(of(

changes(in(populations(and(communities(

Lynn!Govaert1,!Jelena!Pantel1,2!and!Luc!De!Meester1!!1!Laboratory!of!Aquatic!Ecology,!Evolution!and!Conservation,!University!of!Leuven,!Ch.!Deberiotstraat!32,!BL3000!Leuven,!Belgium!2!Centre!d’Ecologie!fonctionelle!et!Evolutive,!Campus!CNRS,!1919!route!de!Mende,!34293!Montpellier!Cedex!5,!France!!During!the!last!decade!there!has!been!an!upsurge!in!interest!to!quantify!the!relative!contribution!of!ecological!and!evolutionary!processes!to!observed!trait!change.!Partitioning!these!contributions!is!a!cornerstone!of!the!field!of!ecoLevolutionary!dynamics.!We!compare!metrics!to!partition!ecological!and!evolutionary!drivers!of!trait!change!that!are!based!on!the!Price!equation!and!on!norms!of!reaction,!and!extend!them!for!the!analysis!of!community!data!and!of!trait!variation!in!space.!We!highlight!advantages!and!limitations!of!the!different!partitioning!metrics!giving!different!results,!with!the!Price!equation!based!partitioning!metrics!not!being!able!to!differentiate!between!genetic!and!nonLgenetic!trait!changes!within!genetic!lineages.!For!data!sets!that!are!both!amenable!to!the!reaction!norm!and!Price!equation!based!partitioning!metrics,!we!advocate!the!use!of!an!extended!metric!that!combines!the!strengths!of!both!approaches.!Applied!at!the!community!level,!this!metric!can!partition!trait!change!in!components!due!to!phenotypic!plasticity!within!genetic!lineages,!genetic!changes!within!lineages,!genetic!changes!due!to!changes!in!frequencies!of!genetic!lineages,!evolution!of!plasticity,!and!components!due!to!changes!in!frequencies!of!species!and!the!gain!and!loss!of!species!in!the!community.!!We!make!a!plea!for!a!careful!choice!of!the!partitioning!metrics!to!be!used!and!take!its!limitations!into!account.!!!!!


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Evolution(of(metamorphosis(in(species(with(an(ontogenetic(diet(shift(

Hanna!ten!Brink1,!André!de!Roos1,!Ulf!Dieckmann2!!1Institute!for!Biodiversity!and!Ecosystem!Dynamics!(IBED),!University!of!Amsterdam,!the!Netherlands!2Evolution!and!Ecology!program,!International!Institute!of!Applied!System!Analysis!(IIASA),!Laxenburg,!Austria!!The!majority!of!all!animal!species!undergoes!a!metamorphosis!at!some!point!during!the!life!cycle.!It!is!thought!that!ontogenetic!changes!in!diet!are!the!first!steps!in!evolutionary!history!towards!life!cycles!with!metamorphosis.!Because!different!resources!require!different!morphologies,!it!is!expected!that!species!with!an!ontogenetic!diet!shift!may!adapt!their!morphologies!over!life!to!the!utilized!resource.!However,!independent!evolution!of!traits!may!be!prevented!by!genetic!correlations!between!the!different!life!stages.!Increasing!fitness!in!one!stage!can!in!this!case!be!disadvantageous!for!another!stage,!leading!to!a!tradeLoff!between!juvenile!and!adult!performance.!It!is!thought!that!metamorphosis!decouples!traits!between!different!lifeLstages,!allowing!the!stages!to!evolve!independently!from!each!other!under!different!selection!regimes.!While!metamorphosis!may!break!up!the!correlations!between!different!life!stages,!it!is!also!a!costly!process.!Species!often!lose!body!mass!during!metamorphosis!and!are!more!vulnerable!to!predation.!This!poses!the!question!under!which!ecological!conditions!the!potential!benefits!of!metamorphosis!can!outweigh!the!disadvantages.!We!used!an!adaptive!dynamics!approach!and!a!sizeLstructured!biomass!model!to!study!which!ecological!conditions!favor!the!evolution!of!metamorphosis!in!species!that!change!their!diet!over!their!life!cycle.!!A(stateOspace(autoregressive(growth(model(to(quantify(individual(heterogeneity(

Mollie!Brooks,!Christopher!Clements,!and!Arpat!Ozgul!!Institute!of!Evolutionary!Biology!and!Environmental!Studies,!University!of!Zurich,!Zurich,!Switzerland!!

Individuals!in!a!population!vary!in!their!growth!rates!due!to!age,!environment,!genetics,!plasticity,!maternal!effects,!cohort!effects,!and!diseases.!As!body!size!has!consequences!for!individual!fitness,!growth!patterns!scale!up!to!affect!population!demography!and!selection.!Here,!we!used!longLterm!data!from!Soay!sheep!of!St.!Kilda!to!quantify!patterns!of!body!mass!growth.!To!enable!the!use!of!nonLconsecutive!observations!and!to!account!for!autocorrelation,!we!fit!the!growth!process!using!a!stateLspace!model.!Our!fixed!effects!included!the!previously!supported!individual!and!environmental!covariates!of!previous!body!mass,!age,!reproductive!status,!population!density,!and!the!North!Atlantic!Oscillation!(NAO)!index!averaged!over!winter.!We!also!included!random!deviates!on!the!intercept!for!each!year,!individual,!and!birth!cohort.!The!individual!random!deviates!represent!among!individual!differences!due!to!unobserved!variables!such!as!genetics!and!parasite!load;!a!positive!deviate!indicates!an!above!average!growth!rate!and!maximum!size.!These!deviates!affect!individuals!throughout!their!ontogeny!and!thus!can!be!used!to!propagate!heterogeneity!in!integral!projection!models!and!ageLstructured!Price!equations!(i.e.!it!is!a!static!rather!than!dynamic!trait).!

We!fit!the!models!using!a!recently!developed!R!package:!TMB.!The!speed!of!TMB!enabled!us!to!do!model!selection!on!2120!versions!of!the!fixed!effect!model!in!less!than!a!day.!We!estimated!the!heritability!of!the!deviates!for!motherLdaughter!pairs,!while!accounting!for!uncertainty.!!

A!model!containing!all!proposed!fixed!effect!terms!except!weight:age2!fell!within!one!deltaAIC!unit!from!the!most!parsimonious!model,!so!we!retained!these!terms.!!We!found!significant!individual!heterogeneity!(CV=0.027,!0.001!SE),!temporal!heterogeneity!(CV=0.012,!0.002!SE),!and!variation!among!cohorts!(CV=0.005,!0.002!SE).!We!found!significant!heritability!(0.32,!0.06!SE)!for!the!growth!deviates!of!mothers!by!daughters.!These!estimates!were!quite!consistent!among!different!fixed!effect!model!versions.!!(

Early(and(extraordinary(physical(peaks(have(a(longevity(cost(

Paul!van!de!Vijver*,!David!van!Bodegom*,!†,!Rudi!GJ!Westendorp*,†,‡!!

*!Leyden!Academy!on!Vitality!and!Ageing,!Rijnsburgerweg!10,!2333!AA!Leiden,!the!Netherlands!†!Leiden!University!Medical!Center,!Albinusdreef!2,!2333!ZA!Leiden,!the!Netherlands!‡!Department!of!public!health,!University!of!Copenhagen,!Nørregade!10,!1017!Copenhagen,!Denmark!(

Introduction(Life!history!theory!postulates!a!tradeLoff!between!investments!in!development!and!longevity.!This!explains!the!divergent!life!courses!of!small,!quick!developing!shortLlived!animals!and!large,!slow!


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developing!longLlived!animals.!We!hypothesized!that!within!humans!those!who!reach!their!physical!peak!early!in!life!or!had!higher!peaks!could!suffer!a!longevity!cost!compared!to!those!who!reach!later!or!lower!physical!peaks.!Material(and(methods(We!used!a!historical!cohort!of!935!Olympic!track!and!field!athletes!born!before!1914!with!a!known!age!at!death.!!Because!of!the!data!on!personal!record!and!the!fact!that!athletes!push!themselves!to!the!limit!we!have!accurate!information!on!the!age!and!magnitude!of!their!physical!peak.!We!standardized!age!and!magnitude!of!personal!record!and!are!therefore!to!be!understood!as!a!relative!age!or!magnitude!compared!to!other!athletes!of!the!same!sex!and!discipline.!Results(In!a!multivariate!analysis!adjusted!for!sex,!nationality,!year!at!birth!and!relative!magnitude,!later!age!at!personal!record!was!accompanied!with!lower!mortality!(HR!0.84!per!SD,!95%!CI!0.77L0.92,!P<0.001).!Higher!magnitude!was!associated!with!higher!mortality!(HR!1.12!per!SD,!95%!CI!1.01L1.24,!P=0.031).!!For!those!whose!personal!record!was!in!the!highest!50%!of!magnitude!and!earliest!50%!of!age!had!a!mean!age!at!death!of!76.7!(SE!1.6)!compared!to!a!mean!age!at!death!of!80.4!(SE!1.3)!in!the!group!with!both!later!age!and!lower!magnitude.!Conclusion(Humans!with!an!early!and!high!physical!peak!have!lower!longevity!than!those!with!a!later!and!lower!physical!peak.!!Part!of!the!variation!in!human!longevity!can!be!explained!by!variation!in!developmental!patterns.!!!Title:(to(be(announced(

James!Holland!Jones,!Stanford!University!(

!Sensitivity(of(the(entropy(of(a(Markov(chain,(or(what(drives(the(rate(of(diversification(in(life(histories(

Uli!Steiner!&!Shripad!Tuljapurkar!!Max!Planck!Odense!Center!on!the!Biodemography!of!Aging!Department!of!Biology,!University!of!Southern!Denmark,!Odense,!Denmark!Department!of!Biology,!Stanford!University,!Stanford,!US!(

The!rate!of!diversification!among!individual!life!courses!can!be!quantified!by!the!entropy!of!the!Markov!chain!that!describes!the!cohort!or!population.!Here!we!present!general!results!for!computing!the!sensitivity!to!this!entropy,!thereby!revealing!drivers!and!evolutionary!forces!that!determine!diversity!in!life!courses!among!individuals.!Some!illustrative!examples!highlight!the!specific!issues!that!arise!particularly!for!stage!structured!populations.!We!discuss!the!findings!with!respect!to!the!evolution!of!demographic!variance!in!life!courses.!!LabyrinthOlike(population(structures(emerge(as(a(consequence(of(multiOlevel(selection(in(selfOorganized(

mussel(beds((

Monique!de!Jager1,!Johan!van!de!Koppel2!&!Franjo!Weissing3!

!1!Ecology!&!Biodiversity!Group,!Institute!of!Environmental!Biology,!Utrecht!University,!Padualaan!8,!3584!CH!Utrecht,!Netherlands.!2!Spatial!Ecology!Group,!Royal!Netherlands!Institute!for!Sea!Research,!Korringaweg!7,!4400!AC!Yerseke,!Netherlands.!!!3!Theoretical!Research!in!Evolutionary!Life!Sciences,!University!of!Groningen,!Nijenborgh!7,!9747!AG!Groningen,!Netherlands.!!

In!selfLorganized!ecosystems,!it!is!eminent!that!groupLlevel!properties!emerge!from!largeLscale!spatial!pattern!formation!that!promote!survival!of!the!organisms!within!the!population.!However,!how!these!emergent!properties!influence!the!evolution!of!selfLorganizing!traits!and!thereby!affect!spatial!pattern!formation!itself!remains!unknown.!Here,!we!demonstrate!that!aggregation!into!clusters!in!selfLorganized!mussel!beds!adds!a!groupLlevel!selection!pressure,!which!can!cause!the!evolution!of!labyrinthLproducing!behaviour!in!mussels.!We!use!a!modelling!approach!that!includes!a!high!amount!of!ecological!detail!to!investigate!the!evolution!of!two!selfLorganizing!traits,!cooperation!and!aggregative!movement,!in!spatially!patterned!mussel!beds,!where!mussels!aggregate!and!attach!byssus!threads!to!neighbouring!conspecifics!in!order!to!decrease!losses!to!predation!and!wave!stress.!We!developed!a!mechanistic,!individualLbased!model!of!spatial!selfLorganization!where!individual!strategies!of!movement!and!attachment!generate!spatial!patterns,!


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which!in!turn!determine!the!fitness!consequences!of!these!strategies.!By!combining!an!individualLbased!simulation!approach!for!studying!spatial!selfLorganization!within!generations!with!an!analytical!adaptive!dynamics!approach!that!studies!selection!pressures!across!generations,!we!are!able!to!predict!how!the!evolutionary!outcome!is!affected!by!environmental!conditions.!When!selection!pressures!on!cooperation!and!movement!are!only!governed!by!local!interactions,!that!is,!the!attachment!of!individuals!to!their!neighbours,!evolution!does!typically!not!result!in!the!labyrinthLlike!spatial!patterns!that!are!characteristic!for!mussel!beds.!However,!when!we!include!a!second!level!of!selection!by!considering!the!additional!protection!provided!by!the!formation!of!mussel!clumps,!evolutionarily!stable!movement!and!attachment!strategies!lead!to!labyrinthLlike!patterns!under!a!wide!range!of!conditions.!!!!!!Physiological(responses(to(light(are(important(predictors(of(growth(in(two(understory(herbs(

Andrea!Westerband,!University!of!Miami,!USA!!

For!plants!in!the!tropical!forest!understory,!the!availability!of!light!strongly!influences!rates!of!growth,!survival,!and!reproduction.!Growth!depends!on!the!successful!conversion!of!carbon!dioxide!to!carbohydrates!via!photosynthesis.!Thus,!demographic!rates!can!be!modelled!as!functions!of!photosynthetic!responses.!I!measured!size,!canopy!openness!(a!proxy!for!light!availability),!and!rates!of!photosynthesis!of!two!locally!coexisting!understory!herbs!in!a!premontane!rainforest!in!Costa!Rica!during!June!2013!and!2014.!Four!physiological!parameters!were!estimated!for!each!individual!using!nonlinear!least!squares.!I!conducted!a!principal!component!analysis!(PCA)!for!both!species,!resulting!in!two!composite!measures!of!physiological!responses.!Model!averaging!was!used!to!predict!future!size!as!a!function!of!current!size,!light!availability,!and!the!composite!physiological!responses.!PCA!revealed!a!high!degree!of!overlap!in!the!grouping!of!physiological!responses!between!the!two!species,!where!the!first!principal!component!axis!was!most!strongly!correlated!with!respiration!and!photosynthetic!capacity!and!the!second!axis!was!most!strongly!correlated!with!photosynthetic!efficiency.!However,!the!average!models!differed!significantly!between!the!species;!current!size!and!light!were!the!most!important!predictors!of!future!size!in!Calathea,!but!current!size!and!photosynthetic!efficiency!were!the!most!important!predictors!of!future!size!in!Heliconia.!Additionally,!both!composite!measures!of!physiology!were!more!important!predictors!of!future!size!in!Heliconia!than!Calathea.!The!twoLway!interaction!between!current!size!and!photosynthetic!efficiency!was!the!most!important!interaction!for!both!species,!but!this!interaction!was!more!important!in!Heliconia!than!Calathea.!In!summary,!physiological!performance!is!a!highly!important!predictor!of!growth!for!plant!species!living!in!resourceLlimited!environments,!and!it!should!be!incorporated!into!future!models!of!population!dynamics.!Such!models!can!provide!insight!regarding!the!physiological!mechanisms!maintaining!species!coexistence.!!Exploring(the(evolution(of(life(history(strategies(using(matrix(population(models(

Owen!Jones1,!Roberto!SalgueroLGomez2!

!1!MaxLPlanck!Odense!Center!on!the!Biodemography!of!Aging!and!Department!of!Biology,!University!of!Southern!Denmark!2!University!of!Queensland,!Australia!!

AgeL!and!stageLbased!matrix!projection!models!(MPMs)!capture!major!components!of!life!history!strategy.!These!strategies!include!the!balances!of!tradeLoffs!among!demographic!components!throughout!the!lifeLcycle!and!mold!mortality!and!fertility!trajectories.!Species’!life!history!strategies!have!been!shaped!by!millions!of!years!of!adaptive!and!neutral!evolution!over!their!shared!phylogenetic!tree.!One!might!expect!closely!related!species!to!share!similar!traits.!Traits!where!this!is!observed!are!said!to!have!high!phylogenetic!signal!but!signal!strength!depends!on!factors!including!the!amount!of!selection!acting!on!the!trait!(i.e.!adaptation),!the!degree!of!environmental!canalization,!and!measurement!error.!We!present!some!results!of!an!exploration!of!phylogenetic!signal!in!demographic!traits!calculated!from!MPMs!in!the!COMPADRE!and!COMADRE!Plant!and!Animal!Matrix!Databases!and!compare!them!to!those!of!morphological!traits.!We!consider!how!our!results!may!affect!the!concept!of!“borrowing!strength”!whereby!unknown!values!for!some!species!can!be!inferred!if!traits!of!relatives!elsewhere!in!the!phylogeny!are!known.!This!is!potentially!of!great!interest!to!workers!in!population!management!and!conservation!fields.!!!!


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!Fertility(decline(in(Japan:(effects(of(socioeconomic(status(and(sexual(conflict(

Masahito!Morita1,2,!Hisashi!Ohtsuki1,!&!Mariko!HiraiwaLHasegawa1!!1Department!of!Evolutionary!Studies!of!Biosystems,!School!of!Advanced!Sciences,!SOKENDAI!(The!Graduate!University!for!Advanced!Studies),!Japan!2Research!Fellow!(DC)!of!the!Japan!Society!for!the!Promotion!of!Science!!

Fertility!directly!affects!one’s!reproductive!fitness,!so!its!decline!is!one!of!the!most!paradoxical!phenomena!in!the!evolution!of!human!behavior.!We!study!the!fertility!decline!in!Japan.!The!fertility!rate!in!Japan!dramatically!dropped!after!the!World!War!II!and!it!has!been!kept!at!a!low!level!despite!the!economic!growth!and!high!resource!availability.!We!study!related!topics!to!fertility!decline!by!using!maltiple!approaches.!

The!current!topics!of!our!studies!are:!(1)!factors!affecting!the!number!of!children!(statistical!analysis!of!crossLsectional!data),!(2)!factors!affecting!the!possibility!of!childbirth!(statistical!analysis!of!panel!data),!and!(3)!sexual!conflict!between!mother!and!father!over!reproductive!decisionLmaking!(questionnaire!survey!at!childcare!facility).!Main!results!of!these!studies!are!as!follows.!(1)’!The!age!at!first!marriage!had!a!significant!effect!on!the!lifetime!number!of!children,!while!one’s!socioeconomic!status,!such!as!income!or!education!level,!did!not.!(2)’!Good!parental!conditions!for!childcare,!such!as!high!income!or!coLresidence!with!parents,!did!not!have!positive!effects!on!the!probability!of!childbirth.!The!presence!of!two!children!strongly!prevented!further!childbirth.!(3)’!We!found!no!sexual!conflict!between!mother!and!father!over!the!ideal!number!of!children.!Also,!each!parent!had!equal!power!to!decide!whether!to!have!children.!

Through!the!aboveLmentioned!studies,!we!have!provided!new!evidence!of!the!preference!in!the!number!of!children.!We!have!also!provided!information!about!sexual!conflict!between!parents!in!reproductive!decisionLmaking!in!a!modern,!developed,!and!lowLfertility!society.!Based!on!these!results,!we!discuss!the!relationship!between!one’s!socioeconomic!success!and!his/her!reproductive!success,!and!the!effects!of!various!kinds!of!sexual!conflict!on!fertility!decline.!!Left(Truncation(in(Human(Survival(Modeling:(Implications(for(Understanding(the(Evolution(of(Heterogeneity(

in(Survival(Capacity(

David!J.!Sharrow!and!James!J.!Anderson!University!of!Washington!!

Early!childhood!and!adult!mortalities,!under!different!evolutionally!forces,!shape!both!the!population!level!and!heterogeneity!of!frailty!or!vitality!within!human!populations.!!Therefore,!linking!the!two!life!stages!as!well!as!clearly!partitioning!the!processes!specific!to!each!is!an!important!task!in!evolutionary!demography.!However,!the!models!to!quantify!the!effects!on!early!life!stages!and!their!contributions!to!adult!mortality!stages!are!very!limited.!As!a!result,!dominant!model!forms!either!left!truncate!to!remove!childhood!mortalities!(e.g.!GompertzLMakeham!model)!or!poorly!fit!the!early!life!patterns!(e.g.!Siler!model).!LeftLtruncation!is!particularly!problematic!because!the!choice!of!truncation!age!strongly!affects!the!estimated!parameters!of!adult!survival.!In!this!paper!we!explore!the!effect!of!left!truncation!age!on!parameter!estimates!in!several!models!and!compare!their!suitability!for!characterizing!the!historical!changes!in!childhood!and!adult!mortality!patterns.!The!implications!for!parameter!interpretations!should!be!of!great!interest!to!researchers!concerned!with!the!biological!and!environmental!influences!on!the!evolution!of!heterogeneity!in!the!survival!capacity!of!humans.!(

Nonlinear(evolutionary(demography(of(complex(life(cycles((

André!de!Roos!Institute!for!Biodiversity!and!Ecosystem!Dynamics,!University!of!Amsterdam!!

A!central!question!in!ecology!and!evolution!is!how!individual!life!history!influences!population!and!community!dynamics!and!in!turn!how!the!population!and!community!setting!influences!selection!and!adaptation!of!life!histories.!Population!impacts!on!individual!life!histories!represent!density!dependence,!which!is!often!limited!to!a!dependence!of!life!history!processes!(e.g.!development,!reproduction,!mortality)!on!total!population!abundance.!Density!dependence,!however,!can!be!represented!far!more!generally!by!distinguishing!the!state!of!the!individual!itself!and!the!environment!it!lives.!Density!dependence!then!emerges!due!to!population!feedback!on!and!thereby!changes!in!the!environment.!!


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Physiologically!structured!population!models!(PSPMs)!are!based!on!this!explicit!recognition!of!the!individual!and!its!environment!and!thereby!consistently!translate!the!individual!life!history!to!the!population!level,!including!the!nonlinear!interactions!individuals!are!exposed!to.!Also,!this!individual/environment!representation!has!formed!the!basis!for!a!general!methodology!that!has!been!developed!in!recent!years!for!demographic,!equilibrium!and!evolutionary!analysis!of!PSPMs.!This!methodology!relies!on!the!numerical!evaluation!of!a!generalised!form!of!the!LotkaLEuler!integral!equation.!Here!I!give!a!brief!introduction!to!the!general!methodology!and!illustrate!how!it!allows!for!evolutionary,!demographic!analysis!starting!from!the!key!ingredients!of!an!individual’s!life!history:!its!development,!reproduction!and!mortality!and!its!interactions,!both!intraL!and!interspecific,!with!its!environment.!!!Variability(of(actuarial(scenescence(patterns(between(captive(and(freeOranging(mammalian(populations(

Tidière!Morgane1,!Gaillard!JM1,!Müller!DWH2,!Gimenez!O3,!Clauss!M4!&!Lemaître!JF1!

1%Université%de%Lyon,%FH69000,%Lyon%;%Université%Lyon%1%;%CNRS,%UMR5558,%Laboratoire%de%Biométrie%et%Biologie%

Evolutive,%FH69622,%Villeurbanne,%France%2%Zoological%Garden%Halle%(Saale),%Fasanenstr.%5a,%06114%Halle%(Saale),%Germany%

3%UMR%5175,%Centre%d’Ecologie%Fonctionnelle%et%Evolutive,%campus%CNRS,%1919%route%de%Mende,%34293,%

Montpellier%Cedex%5,%France%4%Clinic%for%Zoo%Animals,%Exotic%Pets%and%Wildlife,%Vetsuisse%Faculty,%University%of%Zurich,%Winterthurerstr.%260,%

8057%Zurich,%Switzerland%

!Demographic!analyses!based!on!longLterm!studies!have!demonstrated!that!actuarial!(i.e.!survival)!

senescence!is!almost!an!ubiquitous!process!in!vertebrates.!Interestingly!senescence!patterns!can!vary!between!species,!sexes!and!populations.!Among!factors!potentially!explaining!variability!between!populations,!the!most!studied!is!probably!the!environment.!For!instance,!harsh!environments!can!influence!reproductive!effort!during!earlyLlife,!which!will!ultimately!impact!senescence.!

Recently,!it!has!been!shown!that!actuarial!senescence!patterns!differ!between!captive!and!freeLranging!populations!of!large!herbivores.!Generally,!species!have!a!lower!rate!of!senescence!in!captivity!than!in!natural!conditions.!However,!when!the!species!natural!diet!is!complex!(typically!mixed!feeder!species),!the!lower!rate!of!senescence!was!observed!in!freeLranging!conditions!highlighting!the!difficulty!of!keeping!such!browsers!species!in!zoos.!

Here,!we!extended!this!analysis!by!comparing!senescence!patterns!of!captive!and!freeLranging!populations!on!a!wider!range!of!mammalian!species!(from!rodents!to!primates).!In!a!first!step,!we!compared!for!each!species!and!each!sex!the!age!at!the!onset!of!senescence!and!the!rate!of!senescence!between!captive!and!wild!populations.!Then!in!a!second!step,!we!tested!whether!differences!in!senescence!patterns!between!our!two!environmental!conditions!were!explained!by!life!history!traits!(e.g.!position!along!the!slowLfast!continuum)!or!other!ecological!characteristics!(e.g.!natural!diet,!individual’s!home!range!size).!Our!study!emphasize!that!understanding!factors!shaping!betweenLpopulations!differences!in!actuarial!senescence!patterns!is!particularly!important!to!improve!conservation!program!of!protected!species!and!to!upgrade!conditions!in!captivity.!

!Exogenous(and(endogenous(environmental(drivers(of(striped(mouse(demography(

Chloe!Nater1,!Cindy!Canale1,!Koen!van!Benthem1,!Carsten!Schradin2,!Arpat!Ozgul1!%1Institute%of%Evolutionary%Biology%and%Environmental%Studies,%University%of%Zurich%

2Institut%Pluridisciplinaire%Hubert%Curien,%CNRS%Strasbourg%

!Successful!population!management!and!species!conservation!–!especially!in!the!face!of!climate!change!L!

require!detailed!understanding!of!the!impacts!of!environmental!variability!on!ecoLevolutionary!dynamics.!Progressing!towards!this!goal,!longLterm!studies!on!animals!with!large!potential!for!plasticity!and!short!generation!times!may!provide!valuable!insights.!One!such!animal!is!the!African!striped!mouse,!a!small!murid!rodent!which!–!as!a!result!of!evolving!in!an!unpredictable!semiLdesert!–!displays!considerable!phenotypic!plasticity!in!physiology,!behaviour!and!lifeLhistory!traits.!For!this!study,!we!have!used!nine!years!of!markLrecapture!data!from!a!population!in!the!Succulent!Karoo!to!estimate!demographic!rates!as!functions!of!interacting!exogenous!(temperature!and!food!availability)!and!endogenous!(population!density)!environmental!factors!within!a!multiLstate!markLrecapture!framework.!We!then!used!these!functions!in!a!matrix!model!


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approach!to!analyse!the!resulting!populationLlevel!patterns.!We!found!that!complex!interactions!of!environmental!factors!were!responsible!for!variability!in!survival,!whereas!the!regulation!of!reproductive!rates!was!much!simpler.!However,!the!contribution!of!survival!relative!to!reproduction!to!populationLlevel!changes!appeared!to!be!small.!We!conclude!that!population!dynamics!of!our!study!population!are!regulated!mostly!through!changes!in!reproductive!rates!due!to!variability!in!temperature,!food!availability!and!population!density.!We!are!convinced!that!followLup!studies!including!individual!traits,!genetics!and!adaptive!potential!in!addition!to!environmental!drivers!using!the!striped!mouse!study!system!are!well!feasible!and!may!provide!important!contributions!to!our!understanding!of!ecoLevolutionary!dynamics.!!!Consequences(of(stageO(and(sexOspecific(demography(on(avian(adult(sex(ratio(and(beyond(

Luke!J.!EberhartLPhillips1,!Orsolya!Vinzce2,!Cristina!Carmona3,!Clemens!Küpper4,!Joe!Hoffman1,!Tamás!Székely3,!and!Oliver!Krüger1!!

1Department%of%Animal%Behaviour,%Bielefeld%University,%Bielefeld,%Germany%

2Department%of%Evolutionary%Zoology%and%Human%Biology,%University%of%Debrecen,%Debrecen,%Hungary%

3Department%of%Biology%and%Biochemistry,%University%of%Bath,%Bath,%United%Kingdom%

4Institute%of%Zoology,%University%of%Graz,%Graz,%Austria%

!Biased!adult!sex!ratios!in!wild!vertebrate!populations!are!proposed!to!facilitate!the!evolution!of!nonL

traditional!breeding!systems!such!as!polygamy!and!unLparental!care.!!Furthermore,!understanding!the!demographic!consequence!of!adult!sex!ratio!bias!on!population!dynamics!and!viability!is!a!fundamental!step!in!the!applied!fields!of!wildlife!management!and!conservation!biology.!!Here,!we!take!advantage!of!the!highly!tractable!system!provided!by!Charadrius!plovers!to!elucidate!the!stageL!and!sexLspecific!origins!of!biased!adult!sex!ratios.!!Plovers!exemplify!both!interL!and!intraLspecific!variation!in!mating!system!and!parental!care!and!are!demonstrated!to!have!extreme!biases!in!adult!sex!ratio,!thus!providing!an!exciting!group!of!taxa!for!comparative!study!while!controlling!for!phylogenetic!and!ecological!confounds.!!We!used!longLterm!monitoring!data!from!seven!plover!populations!(i.e.,!five!species)!distributed!across!four!continents!to!estimate!sexL!and!stageLspecific!vital!rates!with!markLrecapture!methods,!before!employing!perturbation!analyses!of!twoLsex!matrix!models!to!assess!the!relative!contribution!of!these!vital!rates!on!sex!ratios!of!adult!stable!stage!distributions.!!For!all!seven!plover!populations,!our!modelling!procedure!predicted!sex!ratios!in!the!expected!direction!based!on!published!descriptions!of!breeding!systems.!!For!almost!all!populations,!local!sexLspecific!juvenile!survival!had!the!greatest!influence!on!biasing!adult!sex!ratios,!suggesting!that!sex!differences!in!either!juvenile!dispersal!behaviour!or!juvenile!mortality!potentially!play!important!roles!in!breeding!system!evolution.!!This!finding!raises!exciting!new!questions!regarding!the!flexibility!of!evolutionary!feedbacks!between!population!demography!and!breeding!systems!given!that!juvenile!dispersal!is!a!behavior!mediated!by!an!extrinsic!assessment!of!delayed!future!fitness!prospects,!whereas!juvenile!mortality!is!regulated!by!intrinsic!factors!such!as!genetic!or!physiological!constraints.!!EcoOevolutionary(dynamics(of(populations(dimorphic(for(a(reproductive(trait(–(the(role(of(mutualistic(and(

antagonistic(interactions(

Michel!Thomann1,!Beate!Proske2,!Jon!Ågren1,!Johan!Ehrlén2!!

1!Uppsala!University,!Department!of!Ecology!and!Genetics!!2Stockholm!University,Department!of!Ecology,!Environment!and!Plant!Sciences!!

It!is!well!recognized!that!evolution!can!take!place!over!short!time!scales.!However,!to!what!extent!changes!in!genetic!composition!affect!ecological!processes!and!population!dynamics!is!still!unclear.!!

In!a!field!experiment!we!examined!how!genetic!composition!of!populations!influences!selection!and!population!dynamics!in!the!selfLincompatible!perennial!herb!Primula%farinosa.!On!the!Island!of!Öland!(Sweden)!P.%farinosa!is!dimorphic!for!inflorescence!height!and!natural!populations!are!composed!of!longLscaped!and!shortLscaped!plants!in!different!proportions.!Previous!work!has!shown!that!longLscaped!individuals!are!less!pollenLlimited!but!experience!greater!damage!by!seed!predators!and!large!mammalian!grazers!than!shortLscaped!individuals.!It!has!further!been!shown!that!changes!in!the!grazing!regime!leads!to!changes!in!the!morph!frequencies!of!populations.!

Here!we!examine!if!such!changes!in!genetic!composition!feedback!on!interaction!with!pollinators!and!seed!predators.!To!test!this!we!set!up!a!total!of!25!experimental!populations!with!five!different!morph!frequencies!


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ranging!from!0%!L!100%!longLscaped!individuals!by!altering!the!original!scapeLmorph!composition!of!wild!P.%farinosa!populations.!In!each!population,!flowering!individuals!were!marked!and!grazing!damage,!fruit!set!and!seed!predation!will!be!recorded!at!fruit!maturation!in!2015.!!

We!hypothesize!that!a!higher!proportion!of!the!longLscaped!morph!will!attract!both!more!pollinators!and!predators,!translating!into!an!increased!fruit!set!and!fruit!predation.!As!these!effects!are!operating!in!opposite!directions,!selection!on!scape!morph!and!overall!population!fecundity!will!depend!on!the!relative!strength!of!these!two!effects.!!!Demography(when(history(matters:((A(framework(for(investigating(second(order(matrix(population(models(

Charlotte!de!Vries,!Hal!Caswell!Institute!for!Biodiversity!and!Ecosystem!Dynamics!University!of!Amsterdam!(

In!standard!demographic!models,!the!fate!of!an!individual!is!assumed!to!be!determined!by!the!individual’s!current!condition.!The!validity!of!this!assumption!is!rarely!tested,!with!the!exception!of!a!few!studies!that!found!historical!effects!to!be!important!in!a!plant!species.!!In!this!talk,!we!present!a!general!framework!for!demographic!models!that!incorporate!the!effects!of!history!on!population!dynamics.!The!framework!incorporates!previous!condition!into!the!iLstate!variable!and!includes!an!algorithm!for!constructing!the!population!projection!matrix!from!information!on!current!state!dynamics!as!a!function!of!prior!state.!!This!mathematical!framework!can!be!used!to!analyze!the!sensitivity!of!important!model!outcomes,!like!the!population!growth!rate,!to!historical!effects.!In!other!words,!these!mathematical!tools!can!be!used!to!study!the!effect!of!individual!history!on!individual!fitness.!!!Does(sampling(methodology(affect(population(model(projections(when(there(is(significant(individual(life(

history(variation?(

Anne!M.G.!Kwak1,!Caspar!A.!Hallmann1,!Berend!Voslamber2!&!Eelke!Jongejans1!

!1Radboud!University,!Institute!for!Water!and!Wetland!Research,!Departments!of!Animal!Ecology!and!Ecophysiology!&!Experimental!Plant!Ecology,!PO!Box!9100,!6500!GL!Nijmegen,!The!Netherlands.!2Sovon,!Dutch!Centre!for!Field!Ornithology,!PO!Box!6521,!6503!GA!Nijmegen,!The!Netherlands.!!

Individual!variation!in!life!history!may!arise!through!inherent!differences!between!individuals!(individual!heterogeneity)!or!by!chance!events!(individual!stochasticity).!It!has!been!found!among!many!different!species!and!in!different!elements!of!the!life!cycle!of!those!individuals.!Lifetime!reproductive!success,!for!example,!often!shows!very!large!variation!among!individuals,!both!in!plants!and!animals.!

Even!though!many!studies!have!focused!on!such!individual!variation!in!life!history,!we!found!little!information!on!how!these!differences!affect!population!dynamics!and!population!model!projections.!

Here,!we!use!a!Greylag!Goose!population!(Anser%anser)!as!a!case!study!to!investigate!how!different!sampling!methods!affect!model!outcomes.!Simulated!by!bootstrapping!our!data,!we!create!different!datasets!by:!

1. sampling!over!individuals!their!entire!life!span!(entire!life!histories)!2. sampling!annual!observations!of!individuals!(all!vital!rates!of!an!individual!in!a!specific!year)!3. sampling!annual!observations!per!vital!rate!(i.e.!leaving!out!any!possible!covariance!between!vital!

rates)!The!implications!of!these!sampling!methods!are!explored!with!stochastic!models!and!compared!with!

models!using!the!original!dataset!(no!bootstrapping).!Moreover,!we!applied!a!simulated!dataset!including!a!latent!quality!variable!to!increase!covariance!between!vital!rates.!Using!these!different!sampling!methods!and!datasets,!we!investigate!how!sampling!methodology!affects!population!model!projections,!including!stochastic!population!growth!rate!and!lifetime!reproductive!success.!! !


! 12!

Impacts(of(the(interaction(between(environmental(degradation(and(increasing(environmental(variability(on(

a(population’s(time(to(extinction!Diana!C.!Rypkema,!Meera!Krishnamoorthy,!and!Shripad!Tuljapurkar!Department!of!Biology,!Stanford!University,!Stanford,!CA,!USA!!

The!Earth!is!currently!experiencing!its!sixth!great!mass!extinction.!Previous!mass!extinctions!were!largely!caused!by!climatic!and!ecological!changes,!however!humans!drive!the!current!mass!extinction.!Understanding!the!joint!effects!of!environmental!variability!and!human!disturbance!is!increasingly!important;!future!ecosystems!experiencing!the!strongest!effects!of!climate!change!are!predicted!to!largely!be!postLhuman!disturbance!landscapes.!Although!many!studies!have!looked!at!habitat!degradation!and!increasing!environmental!variability!separately,!few!studies!have!looked!at!the!simultaneous!effect!of!these!two!factors!on!a!population’s!time!to!extinction.!!

We!use!a!simulationLbased!approach!to!find!the!frequency!of!population!extinctions!over!a!variety!of!scenarios!of!habitat!degradation!and/or!increasing!environmental!variability.!We!model!habitat!degradation!by!declining!perLcapita!birth!rates!and!increasing!death!rates.!We!test!each!scenario!over!several!equilibrium!population!sizes!and!life!history!strategies.!

We!found!that!the!combined!force!of!habitat!degradation!and!increasing!environmental!variability!led!to!more!rapid!extinction!than!either!factor!alone.!Specifically,!as!the!doubling!time!of!the!organism!increases,!the!time!to!extinction!also!increases.!We!found!that!for!organisms!with!a!shorter!doubling!time,!habitat!degradation!has!a!significantly!stronger!effect!on!time!to!extinction!than!increasing!environmental!variability.!This!difference!in!relative!importance!of!the!two!factors!underscores!the!significance!of!context!in!our!understanding!of!population!dynamics.!The!framework!developed!here!is!applicable!across!a!broad!range!of!systems!and!can!be!used!to!help!determine!where!to!focus!conservation!and!management!efforts.!!Effects(of(microbial(symbionts(on(host(population(dynamics(in(stochastic(environments(

Teresa!Bohner1!,!Jennifer!Rudgers2!,!Tom!E.X.!Miller3!!

1Dept!of!Plant!Biology,!UCLRiverside,!US,!!2!Dept!of!Biology,!U!New!Mexico,!US,!3Dept!of!BioSciences,!Rice!University,!US!!

Most!multiLcellular!organisms!harbor!microbial!symbionts,!and!many!of!these!are!vertically!transmitted!from!parent!to!offspring.!There!is!growing!awareness!that!microbial!symbionts!can!influence!their!hosts!in!diversity!of!ways,!including!modifications!to!host!physiology,!interactions!with!natural!enemies,!and!responses!to!environmental!stress.!These!effects!can!vary!in!magnitude!and!direction,!though!vertical!transmission!is!generally!expected!to!favor!the!evolution!of!hostLsymbiont!mutualism!(positive!effects).!While!the!individualLlevel!effects!of!microbial!symbionts!on!host!organisms!are!increasingly!well!documented,!relatively!little!is!known!about!their!role!in!host!population!dynamics.!This!is!especially!true!for!host!populations!in!stochastic,!randomly!varying!environments!(i.e.,!most!natural!environments).!Theory!suggests!that,!in!a!stochastic!environment,!symbionts!could!affect!host!population!dynamics!in!either!or!both!of!two!ways:!by!modifying!their!mean!vital!rates!and!/!or!the!temporal!variance!in!vital!rates.!No!empirical!studies,!as!far!as!we!are!aware,!have!attempted!to!quantify!this!influence!and!decompose!the!relative!importance!of!effects!on!vital!rate!means!vs!variances.!To!fill!this!gap,!we!combined!field!experiments!and!demographic!models!to!study!the!heritable!symbiosis!between!a!perennial!coolLseason!grass!(Poa%autumnalis)!and!its!vertically!transmitted!fungal!endophyte!(Epichloe!sp.).!We!created!experimental!populations!of!plants!that!harbored!the!endophyte!(E+)!and!plants!from!which!endophytes!were!experimentally!removed!(EL).!We!tracked!demographic!vital!rates!in!these!experimental!populations!over!eight!years.!With!hierarchical!Bayesian!models,!we!found!that!endophytes!modify!both!the!means!and!the!temporal!variances!of!most!vital!rates.!We!used!Life!Table!Response!Experiments!to!quantify!the!extent!to!which!the!net!effect!of!symbiosis!was!due!to!effects!on!means!vs!variances.!Our!results!suggest!that!heritable!symbionts!could!have!important!effects!on!host!population!dynamics!by!buffering!against!negative!effects!of!temporal!variability.!This!new!finding!that!suggests!that!important!effects!of!microbial!symbioses!on!host!fitness!and!population!dynamics!can!only!be!observed!in!the!context!of!longLterm!studies!that!include!realistic!environmental!variability.!!!!!!


! 13!

The(COMADRE(Animal(Matrix(Database:(an(open(resource(for(comparative(animal(demography(

Roberto!SalgueroLGómez1,2,!Owen!R.!Jones3,4,!C.!Ruth!Archer1,5,!Yvonne!M.!Buckley2,7,!Judy!CheLCastaldo8,!David!Hodgson9,!Alexander!Scheuerlein1,!Hal!Caswell10,11,!&!James!W.!Vaupel1,3,12!!

1!Laboratory!of!Evolutionary!Biodemography!Laboratory,!Max!Planck!Institute!for!Demographic!Research.!KonradLZuseLStraße!1,!Rostock,!DEL18057,!Germany.!2!The!University!of!Queensland,!Australia.!3!MaxLPlanck!Odense!Center!on!the!Biodemography!of!Aging,!University!of!Southern!Denmark!4!Department!of!Biology,!University!of!Southern!Denmark.!5!Max!Planck!Institute!for!Demographic!Research,!Rostock,!Germany.!6!Department!of!Plant!Science!and!Landscape!Architectur,!University!of!Maryland.!7!School!of!Natural!Sciences,!Trinity!College!Dublin.!8!National!SocioLEnvironmental!Synthesis!Center.!Annapolis,!MD,!USA.!9!Centre!for!Ecology!and!Conservation,!University!of!Exeter,!UK.!10!Institute!for!Biodiversity!and!Ecosystem!Dynamics,!University!of!Amsterdam!11Biology!Department,!Woods!Hole!Oceanographic!Institution.!12!Population!Research!Institute.!Duke!University.!!

Matrix!population!models!(MPMs)!are!among!the!most!widely!used!demographic!tools!by!animal!ecologists.!MPMs!project!population!dynamics!in!terms!of!reproduction,!mortality,!and!development!over!the!entire!life!cycle.!The!results!of!MPMs!have!direct!biological!interpretations,!facilitating!comparisons!among!animal!species!as!different!as!Caenorhabditis%elegans,!Loxodonta!africana!and!Homo%sapiens.!Thousand!of!animal!demographic!records!exist!in!the!form!of!MPMs,!but!they!are!dispersed!throughout!the!literature,!rendering!comparative!analyses!difficult.!Here,!we!introduce!the!COMADRE!Animal!Matrix!Database,!an!openLsource!online!repository,!which!in!its!version!1.0.0!contains!data!on!346!species!worldwide,!from!280!studies!with!a!total!of!1,603!population!projection!matrices.!COMADRE!also!contains!ancillary!information!(e.g.!ecoregion,!taxonomy,!biogeography,!etc.)!that!facilitates!interpretation!of!the!numerous!demographic!metrics!that!can!be!derived!from!its!MPMs.!The!open!access!nature!of!this!database,!together!with!its!ancillary!information!will!facilitate!comparative!analysis,!as!will!the!growing!availability!of!databases!focusing!on!other!traits,!and!tools!to!query!and!combine!them.!Through!future!frequent!updates!of!COMADRE,!and!its!integration!with!other!online!resources,!we!showcase!some!of!the!analyses!on!comparative!demography!and!life!history!theory!that!can!be!undertaken!based!on!COMADRE!and!its!sister!database,!the!COMPADRE!Plant!Matrix!Database.! !!

Tuesday,(6(October(2015(

! 14!

!!

Tuesday!6th!of!October!2015!

! Presenter!! Title!(abridged)! !09:00L09:30!!

Keynote%4:!Oskar!Burger! On!the!pressing!need!for!more!evolutionary!demography!of!human!fertility!and!population!growth!

09:30L09:40! Marjolein!Bruijning! Using!automated!particle!tracking!...!09:40L09:50! Michela!Busana! Demographic!consequences!of!cooperative!behavior!...!09:50L10:00! Callum!Lawson! Are!betLhedgers!better!prepared!for!climate!change?!10:00L10:10! Jennifer!Williams! Plants!on!the!move:!roles!of!evolution!and!landscape!structure...!10:10L10:20! Lalasia!BialicLMurphy! FineLscale!habitat!heterogeneity!...!10:20L10:30! Carol!Horvitz!! Spatial!variability!in!transient!elasticity!...!10:30L11:00!!

Coffee(break(

11:00L11:10! Susanne!Schindler! SexLspecific!demography!and!TriversLWillard!!11:10L11:20! Lionel!Jouvet! Lifespan!and!reproduction!of!isoclonal!individual!E.%coli...!11:20L11:30( Koen!van!Benthem! Disentangling!evolutionary!and!plastic!processes!...!11:30L11:40! Merel!Jansen! Is!the!tradeLoff!between!growth!and!tolerance!...!heritable?!11:40L11:50! Ilona!Nenko! Relationship!between!reproductive!effort!and!cognitive!function!11:50L12:00! Heike!Gruber! Environmental!impact!on!lifeLhistory!strategies...!12:00L12:30! Keynote%5:!Samuel!Pavard! How!cognition!and!sociality!have!shaped!human!biology:!a!

biodemographic!perspective!12:30L13:30!!

Lunch(

13:30L14:00!!

Keynote%6:!Hans!Metz! Evolutionary!demography!

14:00L14:10! Nora!Sánchez!Gassen! Sensitivity!of!population!growth!rate!...!transient!and!asymptotic!14:10L14:20! JeanLFrançois!Lemaître! Mother's!curse!and!the!evolution!of!sexLspecific!patterns...!14:20L14:30! Lin!Chen! Risk!perception's!influences!on!teenage!childbearing...!14:30L14:40! Tina!Cornioley! Wind!is!an!important!climatic!factor!...!14:40L14:50! Yngvild!Vindenes! Evaluating!the!fitness!importance!of!maternal!size!effects!...!14:50L15:00! Rebecca!Sear! Consequences!of!early!life!stress!for!fertility!...!15:00L15:30!!

Keynote%7:!Jacob!Moorad! Evolution!of!maternal!effect!senescence!

15:30L16:00!!

Coffee(break(

16:00L16:10! Eelke!Jongejans! DemographyLbased!assessment!of!bird!population!trends!16:10L16:20! Kate!LaytonLMatthews! Environmental!drivers!of!metapopulation!dynamics...!16:20L16:30! Gregory!Roth! Introducing!hyperLmatrix!models!16:30L16:40! Felix!Tropf! MegaLanalysis!of!31,396!individuals!beyond!reproductive!age...!16:40L16:50! Orou!Gaoue! Climate,!not!harvest,!is!the!main!driver!of!ageLspecific!mortality...!16:50L7:20! Keynote%8:!Tiffany!Knight!

!Consequences!of!pollen!limitation!for!plant!population!dynamics!

17:20L18:20! Drinks(+(posters(

18:30! Dinner(

!!!!! !


! 15!

On(the(pressing(need(for(more(evolutionary(demography(of(human(fertility(and(population(growth(

!1. Oskar!Burger,[email protected],!University!of!Kent,!School!of!Anthropology!and!Conservation,!

Canterbury,!UK!and!Max!Planck!Institute!of!Demographic!Research,!Rostock,!Germany!!2. John!DeLong,!School!of!Biological!Sciences,!University!of!Nebraska,!Lincoln,!USA!3. Rebecca!Sear,!London!School!of!Hygiene!and!Tropical!Medicine,!London,!UK!!!

Evolutionary!demographers!have!made!great!strides!in!applying!evolutionary!principles!to!human!phenomena!in!spite!of!a!sometimes!hostile!reception!from!social!scientists.!However,!much!of!the!progress!in!evolutionary!demography!has!come!from!studies!of!mortality!and!ageing,!which!are!somewhat!less!controversial!because!they!rely!less!on!explanations!of!cognition!or!behaviour.!For!this!reason,!human!fertility!tends!to!be!the!focus!of!mainstream!demography,!economics,!and!anthropology,!but!is!lessLstudied!by!evolutionary!scientists.!We!argue!that!much!more!comparative!and!theoryLdriven!work!is!needed!on!the!topic!of!fertility!variation!within!and!among!human!populations,!including!the!challenge!of!the!radical!declines!to!belowLreplacement!fertility!in!the!postLindustrial!age.!We!highlight!key!areas!of!overlap!between!evolutionary!and!nonLevolutionary!approaches!to!fertility!variation!and!emphasize!key!areas!where!the!two!approaches!are!at!odds.!We!are!particularly!sceptical!of!the!assumption!in!“demographic!transition!theory”!that!fertility!decline!is!irreversible.!We!present!theoretical!rationales!for!plasticity!in!fertility!and!review!pathways!by!which!fertility!might!not!be!expected!to!permanently!decline!globally!to!one!fixed!number.!The!assumption!of!irreversibility!is!critical!as!it!guides!policy!and!population!projection!in!many!ways.!While!human!fertility!behaviours!may!be!especially!complex!the!combined!toolkits!of!demography!and!evolutionary!ecology!can!shed!novel!insight!on!the!problem!of!very!low!fertility.!!

!!

Using&automated&particle&tracking&to&obtain&demographic&information&from&video&sequences&in&R!

Marjolein*Bruijning,"Marco"D."Visser,"Eelke"Jongejans!Radboud'University,'Nijmegen!

Many%demographic%studies,%both%experimental%and%observational,%require%temporal%data%on%population%counts'and'body'size'distributions.'Until'now,'measurements'have'largely'been'done'manually,'putting'a'limit'on#the#amount#of#replicates,#study#objects#or#densities&that&can&be&treated.&Although&particle&identification&and&tracking)algorithms)do)exist)in)other)fields)of)research,)they)have)hardly)been)applied)by)demographers)and)ecologists.*We*have*developed*a*general*and*straightforward*method*to*obtain*estimates%of%population%densities'and'individual'body'sizes,'using'the'statistical'programming'language'R.'The'method,'which'uses'video&footage&as&input,&is&based&on&background&subtraction,&thresholding&and&connected&component&labeling&to&detect%moving%particles.#This#is#done#in#combination#with#a#trained#artificial#neural#network,#to#optimize#detection(and(noise(filtering.(After(identifying(all(particles,(a(tracking(algorithm(records(individual(paths,(based(on#distance#between#particles,#movement#history#and#particle'size.'This'yields'information'on'movement'behavior,*speed*and*body*size*distributions.*In*addition,*when*using*a*time*series*of*video*material,*our*method'can'directly'provide'estimates'of'survival,'growth'and'reproduction,'based'on'shifts'in'densities$and$body%size%distributions.%We%have%successfully%applied%the%developed%method%on%populations%of%Daphnia!magna%in#a#laboratory#setting.#In#this#setting,#our#method#enabled#fast#and#accurate#estimates.#We#plan#to#test#the#applicability)and)accuracy)on)other%systems,%including%other%aquatic%laboratory%systems%and%insect%populations%in#field#situations.!!Demographic(Consequences(of(Cooperative(Behavior(

Michela!Busana1*,!Dylan!Childs2,!Jan!Komdeur1,!Margarete!Utz1,!Terry!Burke2,!David!Richardson3!and!Hannah!Dugdale1,2!!

1!Groningen!Institute!for!Evolutionary!Life!Sciences,!University!of!Groningen,!Groningen,!NL!2!Department!of!Animal!and!Plant!Sciences,!University!of!Sheffield,!Sheffield,!UK!3!School!of!Biological!Sciences,!University!of!East!Anglia,!UK!!

Cooperative!behavior!is!often!defined!as!a!social!interaction!when!more!than!2!individuals!live!in!a!group!and!provide!care!for!the!young!of!a!single!brood!or!litter.!These!groups!are!usually!characterized!by!strong!


! 16!

social!structures!with!dominant!breeders!and!subordinates.!Subordinates!delay!their!reproduction!and!help!the!dominants!with!the!care!of!their!young.!The!dominance!position!in!the!social!structure!is!often!associated!with!variations!in!morphological!traits.!Both!the!dominance!position!and!morphological!traits!correlate!with!variations!in!individual!fitness.!Understanding!the!relationships!between!dominance!statuses,!individual!traits!and!fitness!helps!define!the!longLterm!demographic!consequences!of!cooperation!on!population!dynamics.!Reliable!population!projections!are!needed!to!manage!populations,!especially!when!they!are!of!conservation!concern.!

Seychelles!warblers!are!endemic!passerines!living!on!5!islands!in!the!Seychelles!archipelago!and!are!listed!as!vulnerable!in!the!IUCN!Red!List.!They!are!facultative!cooperative!breeders.!Their!groups!are!constituted!by!the!dominant!breeding!pair!and!up!to!4!subordinates!(average!group!size!=!3.3!individuals,!SD!=!1.2).!Some!of!these!subordinates!may!help!the!dominant!pair!raise!young,!while!others!reside!within!the!group,!but!do!not!provide!help.!Previous!studies!showed!that!population!density!is!negatively!related!with!reproductive!rates,!but!there!is!no!densityLdependence!effect!on!individual!survival.!We!applied!an!Integral!Projection!Model!to!the!Seychelles!warbler!to!test!for!the!simultaneous!effect!of!individual!body!sizes,!dominance!status,!density!dependence!and!helping!behavior!on!population!stability.!We!discuss!how!the!presence!of!helping!behavior!buffers!against!fluctuations!in!population!size!and!influences!the!distribution!of!individual!traits!in!the!population.!Despite!the!widespread!presence!of!cooperative!behavior!(~3%!of!birds!and!mammals!are!cooperative);!the!application!of!individual!based!demographic!models!remains!limited.!Including!individual!variations!in!body!size!improves!the!ability!to!make!accurate!demographic!predictions!of!cooperatively!breeding!populations.!

*!Corresponding!author.!ELmail:[email protected]!!!Are(betOhedgers(betterOprepared(for(climate(change?(

Callum!Lawson1,!D.!Lawrence!Venable2,!Jennifer!Gremer3!&!Martijn!van!de!Pol1,4!

(1Department%of%Animal%Ecology,%Netherlands%Institute%of%Ecology%(NIOOHKNAW)%

2Department%of%Ecology%and%Evolutionary%Biology,%University%of%Arizona,%Tucson,%USA%

3US%Geological%Survey,%Southwest%Biological%Science%Center,%Flagstaff,%USA%

4Division%of%Evolution,%Ecology%&%Genetics,%The%Australian%National%University%

%

Responses!to!climate!change!vary!greatly!among!species,!due!to!differences!in!key!life!history!traits!such!as!body!size,!offspring!numbers,!or!development!rates.!These!traits!have!been!shaped!by!past!environmental!conditions:!for!example,!species!exposed!to!greater!temporal!environmental!fluctuations!have!tended!to!evolve!“betLhedging”!life!history!traits,!such!as!development!rates!that!vary!among!individuals,!which!prime!them!to!survive!unfavourable!periods.!In!addition!to!being!adapted!to!past!environmental!variation,!species!with!these!betLhedging!lifeLhistories!may!be!betterLequipped!to!cope!with!the!changes!in!the!means!and!variances!of!environmental!conditions!that!are!occurring!under!climate!change,!but!this!prediction!has!rarely!been!tested!empirically.!In!this!talk,!I!use!longLterm!data!on!the!population!dynamics!of!desert!annual!plants!to!assess!whether!species!with!betLhedging!adaptations!are!more!resistant!to!climate!change.!Specifically,!I!test!whether!species!with!higher!levels!of!seed!dormancy!(an!adaptation!to!interLannual!variance!in!rainfall)!will!have!higher!persistence!under!increasingly!variable!and!arid!conditions.!The!results!provide!an!empirical!test!of!whether!betLhedging!lifeLhistory!traits!can!be!used!to!identify!species’!vulnerabilities!to!climate!change.!

!!

Plants(on(the(move:(roles(of(evolution(and(landscape(structure(in(spreading(populations(

Jennifer!L.!Williams!(University!of!British!Columbia)!&!Jonathan!M.!Levine!(ETH!Zurich)!!

Predicting!the!spatial!advance!of!biological!invasions!and!rangeLshifting!native!species!under!climate!change!requires!understanding!how!evolutionary!processes!influence!the!velocity!of!spread.!Although!landscape!heterogeneity!is!known!to!influence!the!ecological!dynamics!of!spreading!populations,!its!effect!on!evolutionary!dynamics!influencing!spread!is!poorly!understood.!We!explored!how!landscape!heterogeneity!changes!selection!in!advancing!populations!and!the!degree!to!which!evolution!can!accelerate!invasion!velocities!using!the!experimental!invasions!of!the!annual!plant!Arabidopsis%thaliana.!In!the!greenhouse,!we!invaded!experimental!‘runways’!with!multiple!genotypes!of!Arabidopsis!populations.!We!compared!how!invasion!velocity!differs!between!treatments!with!evolution!and!where!evolution!is!effectively!stopped!using!a!


! 17!

treatment!to!reset!each!generation!back!to!the!initial!seeding!frequencies,!and!across!gap!sizes.!After!five!generations!of!spread,!we!found!that!the!invasion!velocity!was!significantly!higher!when!evolution!was!acting,!and!that!this!increase!was!greater!for!landscapes!with!larger!gap!sizes.!This!result!is!in!contrast!to!our!recent!theoretical!work,!which!found!that!although!evolutionary!change!could!increase!the!spread!velocity,!this!increase!could!be!insignificant!in!patchy!landscapes,!where!competitive!traits!were!favored!in!both!spreading!and!nonLspreading!populations.!Finally,!we!found!that!some!genotypes!have!become!more!common!and!dominate!the!invasion!fronts!compared!to!the!population!cores.!We!will!assess!which!traits!are!associated!with!these!genotypes!and!compare!them!to!the!theoretical!predictions!across!landscape!types.!!!FineOscale(habitat(heterogeneity(and(a(nonOnative(avian(frugivore(drive(the(population(dynamics(of(an(island(

endemic(shrub,(Cyrtandra)dentata)%

Lalasia!BialicLMurphy1,!Orou!G.!Gaoue1,2!!1!Department!of!Botany,!University!of!Hawai‘i!at!Manoa!3190!Maile!Way,!St.!John!101,!Honolulu,!Hawai‘i!96822,!USA;!2!Universite!de!Parakou,!BP!123,!Parakou,!Benin.!(

For!endangered!plants!with!small!population!size,!fineLscale!habitat!heterogeneity!can!limit!population!persistence.!On!islands,!seed!predation!by!nonLnative!animals!can!drastically!reduce!the!reproductive!success!of!native!plants.!In!this!study,!we!examined!the!combined!influence!of!a!nonLnative!avian!frugivore,!redLbilled!leiothrix!(Leiothrix%lutea),!and!fineLscale!habitat!heterogeneity!on!the!population!dynamics!of!an!island!endemic!longLlived!shrub,!Cyrtandra%dentata.!At!the!field!site,!redLbilled!leiothrix!reduced!seed!rain.!Furthermore,!fineLscale!habitat!heterogeneity!strongly!correlated!with!the!spatial!distribution!of!C.%dentata.!Specifically,!seedling!establishment!was!significantly!greater!on!talus!rocky!outcrops!covered!by!moss,!relative!to!soil.!The!interaction!of!C.%dentata!with!the!removal!of!redLbilled!leiothrix!and!the!loss!of!optimal!safeLsites!for!seedling!establishment!influenced!the!short!and!longLterm!population!dynamics.!Although!the!individual!interaction!of!C.%dentata!with!these!environmental!stressors!had!a!negative!influence!on!population!dynamics,!they!did!not!change!the!direction!of!the!population!growth!rate!(e.g.,!growing!or!declining).!However,!the!combined!impact!of!redLbilled!leiothrix!and!the!loss!of!optimal!habitat!for!seedling!establishment!caused!the!population!growth!rate!to!shift!from!growing!to!declining.!Additionally,!the!life!stage!and!associated!vital!rate!that,!if!perturbed,!had!the!greatest!relative!impact!on!the!population!growth!rate!was!the!survival!of!mature!plants,!followed!by!the!growth!of!seedlings!and!immature!plants.!The!results!of!this!study!emphasize!the!need!to!unravel!the!synergistic!effect!of!environmental!stressors!when!planning!and!prioritizing!rare!plant!restoration.!!!Spatial(variability(in(transient(elasticity:(implication(for(evolutionary(ecology(of(novel(interactions(and(

application(to(biocontrol(of(an(invasive(plant(in(Hawaii(

!Carol!C.!Horvitz,!Department!of!Biology,!Institute!of!Theoretical!and!Mathematical!Ecology,!University!of!Miami,!PO!Box!249118,!Coral!Gables,!FL!33124,!USA;!Julie!S.!Denslow,!Department!of!Biology!and!Evolutionary!Biology,!Tulane!University,!New!Orleans,!LA!70130,USA;!!Orou!Gaoue,!University!of!Hawaii!at!Manoa,!Department!of!Botany,!3190!Maile!Way,!St.!John!101,!Office!405C,!Honolulu,!HI!96822,!USA;!Tracy!Johnson,%Institute!of!Pacific!Islands!Forestry,!USDA!Forest!Service,!Volcano,!HI!96785,USA!!

The!goal!of!bioLcontrol!is!to!reduce!populations!of!targeted!pests!over!a!large!region.!!An!important!first!step!is!to!obtain!and!analyze!preLrelease!demographic!data!on!the!pest!at!more!than!one!site.!Here,!we!focus!on!short!term!dynamics,!which!will!be!operating!at!the!time!a!biocontrol!agent!is!released.!Transients!differ!from!asymptotic!dynamics!most!when!(1)!the!stage!structure!is!far!from!the!asymptotic!stage!structure!(this!distance!weighted!by!reproductive!value!is!measured!by!α0)!and!(2)!the!nonLdominant!dynamics!are!strong!relative!to!the!dominant!(measured!by!the!damping!ratio,!ρ).!To!determine!site!differences!in!the!expected!effects!of!an!insect!bioLcontrol!agent!(Tectococcus%ovatus)!on!populations!of!the!invasive!pest!plant!Psidium%

cattleianum!(Strawberry!guava)!in!Hawaii,!we!use!detailed,!demographic!data!collected!over!multiple!preLrelease!years,!to!parameterize!an!integral!projection!model!of!population!dynamics!at!four!sites.!!We!found!preLrelease!transient!dynamics!varied!across!the!sites!in!both!α0!and!ρ!as!well!as!in!the!elasticity!of!short!term!growth!to!different!types!of!demographic!rates!(growth!of!seedlings,!growth!of!adults,!reproduction!by!seed!


! 18!

and!reproduction!by!vegetative!sprouting).!!The!initial!stage!distributions!differed!from!their!siteLspecific!asymptotic!distributions!in!different!ways.!Two!sites!started!with!excess!individuals!in!high!RV!stages,!α0!>0;!while!two!sites!start!with!excess!individuals!in!low!RV!stages,!α0!<0.!The!importance!of!nonLdominant!dynamics,!ρ,!ranged!!from!0.79!to!0.99;!the!site!with!the!largest!value!is!the!one!where!transients!will!remain!most!dissimilar!from!asymptotics!for!a!longer!period!of!time.!The!changing!stage!distribution!of!a!population!during!the!transient!phase!coupled!with!analytical!transient!elasticities!drive!these!changes,!with!implications!for!evolutionary!ecology!of!novel!interactions!in!general.!!!SexOspecific(demography(and(TriversOWillard(

Susanne!Schindler,!University!of!Zurich!(Switzerland)!!and!University!of!Oxford!(UK)!!JeanLMichel!Gaillard,!Universite!Claude!Bernard!Lyon!1!(France)!Andre!Gruening,!University!of!Surrey!(UK)!Peter!Neuhaus,!!University!of!Calgary!(Canada)!Lochran!W.!Traill,!University!of!the!Witwatersrand!(South!Africa)!Shripad!Tuljapurkar,!Stanford!University,!Stanford!(USA)!Tim!Coulson,!University!of!Oxford!(UK)!

!TriversLWillard!theory!proposes!that!the!sex!ratio!of!offspring!should!vary!with!maternal!condition!when!it!has!sexLspecific!influences!on!offspring!fitness.!In!particular,!mothers!in!good!condition!in!polygynous!and!dimorphic!species!are!predicted!to!produce!an!excess!of!sons,!whereas!mothers!in!poor!condition!should!do!the!opposite.!Despite!the!elegance!of!the!theory,!support!for!it!has!been!limited.!Here,!we!extend!and!generalise!the!TriversLWillard!theory!to!help!explain!the!mismatch!between!predictions!and!observations!of!offspring!sex!ratio.!In!polygynous!species,!males!typically!have!higher!mortality!rates,!different!ageLspecific!reproductive!schedules,!and!more!riskLprone!life!history!tactics!than!females.!!These!differences!are!not!currently!incorporated!into!TriversLWillard!theory.!Using!twoLsex!models!parameterised!with!data!from!freeLliving!mammal!populations!with!contrasting!levels!of!sexLdifferences!in!demography!we!demonstrate!how!sexLdifferences!in!life!history!traits!over!the!entire!lifespan!can!lead!to!a!wide!range!of!sex!allocation!tactics.!Correlations!between!maternal!condition!and!offspring!sex!ratio!!alone!are!insufficient!to!conclude!that!mothers!are!adaptively!adjusting!offspring!sex!ratio.!!Lifespan(and(reproduction(of(isoclonal(individual(E.coli(in(different(environments!Lionel!Jouvet!&!Ulrich!Steiner!Max!Planck!Odense!Center!on!the!Biodemography!of!Aging!&!Department!of!Biology,!University!of!Southern!Denmark,!Campusvej!55,!Odense,!Denmark!!Email:[email protected]!!Lifespan!and!reproduction!are!key!fitness!components,!both!of!which!are!influences!by!genetics!and!the!environment.!Tracking!large!numbers!of!genotypically!known!individuals!throughout!their!lives!in!known!environments!has!been!challenging.!Here!we!show!for!isogenic!individual!E.!coli!bacteria!under!controlled!environments!how!demographic!parameters!and!distributions!in!reproduction!and!survival!change!across!environments.!We!achieve!this!by!using!a!microfluidic!device!that!traps!thousands!of!individual!E.!coli!cells!and!tracks!their!division!(reproduction)!over!their!lifespan.!Our!results!show!substantial!variation!of!lifespan!and!reproduction!within!the!same!environment,!and!significant!shifts!in!mean!and!variance!in!lifespan!and!reproduction!among!environments.!Both!variance!and!mean!lifespan!and!reproduction!decreases!with!increased!temperature,!but!the!ratio!between!the!two!remains!surprisingly!constant.!The!environmental!dependency!of!neutral!variability!among!individuals!suggests!different!adaptive!paces!across!environments.!! !


! 19!

Disentangling(evolutionary(and(plastic(processes(underlying(trait(dynamics(

!Koen!van!Benthem1!,!Marjolein!Bruijning2!,!Timothée!Bonnet1!,!Eelke!Jongejans2!,!Erik!Postma1,!!Arpat!Ozgul!1!!1Institute!of!Evolutionary!Biology!and!Environmental!Studies,!University!of!Zurich!(CH)!2Department!of!Animal!Ecology!and!Ecophysiology,!Radboud!University!(NL)!!

Multiple!frameworks!have!been!developed!that!allow!disentangling!the!evolutionary!and!ecological!processes!(including!selection,!plasticity!and!demography)!underlying!phenotypic!change!over!ecological!timespans.!!Doing!so,!generates!a!better!understanding!of!trait!dynamics!in!natural!populations.!However,!their!(simultaneous)!application!to!empirical!datasets!remains!scarce,!and!hence!it!remains!unclear!how!they!compare!to!each!other.!Here!we!focus!on!four!main!frameworks:!the!quantitative!genetic!animal!model,!the!genotypeLphenotypeLenvironment!equation,!the!ageLstructured!Price!equation!and!integral!projection!models.!By!applying!each!of!these!to!the!same!individualLbased!and!genetically!explicit!simulated!dataset,!we!show!that!these!frameworks!come!to!different!conclusions!with!respect!to!which!processes!contribute!most!to!trait!dynamics.!This!is!due!to!differences!in!the!estimation!procedure,!as!well!as!to!the!way!the!different!processes!are!defined!in!each!framework.!Furthermore,!we!investigate!the!theoretical!limits!of!the!frameworks.!On!the!whole,!this!work!will!aid!the!interpretation!of!the!results!from!each!of!these!frameworks,!and!provides!insight!into!which!framework!suits!which!biological!question!best.!!Is(the(tradeOoff(between(growth(and(tolerance(to(herbivory(heritable?(A(test(for(an(understorey(palm(

!Merel!Jansen1,2,3,!Pieter!Zuidema2,!Frans!Bongers2,!Miguel!MartínezLRamos3,!Aad!van!Ast1,!Niels!Anten1!1%Centre%for%Crop%Systems%Analysis,%Wageningen%University,%P.O.%Box%430,%6700%AK,%Wageningen,%The%

Netherlands%2Forest%Ecology%and%Forest%Management%group,%Wageningen%University%

3%Centro%de%Investigaciones%en%Ecosistemas,%Universidad%Nacional%Autónoma%de%México%

!Herbivory!is!an!ubiquitous!stressor!that!can!strongly!limit!individual!performance.!The!most!studied!

mechanisms!by!which!plants!counteract!this!involve!plant!defences!(plant!resistance)!that!(partially)!prevent!herbivory.!But!over!the!last!decades!it!has!become!clear!that!many!species!have!evolved!tolerance;!i.e.,!the!ability!of!plants!to!maintain!performance!when!suffering!leaf!loss,!as!a!strategy!to!mitigate!the!negative!effects!of!herbivory,!e.g.!by!investment!in!reserves.!However,!as!energy!can!only!be!used!once,!allocational!tradeLoffs!may!exist!between!tolerance!to!herbivory!and!other!characteristics!such!as!growth!rate,!which!are!reflected!in!different!life!histories.!The!tradeLoff!between!tolerance!and!growth!has!been!shown!to!exist!between!many!species,!but!for!this!to!have!evolved,!in!theory!it!should!also!exist!within!species.!However,!so!far!very!little!is!known!about!the!existence,!heritability!and!strength!of!toleranceLgrowth!tradeLoffs!within!species!and!the!extent!to!which!this!can!lead!to!withinLpopulation!variation!in!life!histories.!

In!this!study!we!analysed!to!what!extent!growth!and!tolerance!to!defoliation!are!heritable!and!if!a!tradeLoff!between!the!two!exists.!We!did!this!for!the!shade!tolerant!understorey!palm!Chamaedorea%elegans,!using!data!from!a!quantitative!genetic!greenhouse!experiment!with!1440!seedlings!from!202!halfLsib!families,!where!seeds!where!obtained!from!a!natural!population!in!Mexico.!A!2/3!defoliation!treatment!was!applied!to!half!of!the!individuals!to!simulate!herbivory.!We!found!that!growth!rate!was!strongly!heritable,!and!that!slow!growing!families!where!indeed!more!tolerant!to!defoliation!in!terms!of!growth.!Furthermore,!we!constructed!a!demographic!model!based!on!data!from!a!long!term!field!experiment,!in!which!we!incorporated!heritability!estimates!of!growth!and!tolerance.!With!this!model!we!analysed!to!what!extent!the!found!tradeLoff!can!lead!to!heterogeneity!in!life!histories.!!!Is(there(a(relationship(between(reproductive(effort(and(cognitive(function(in(postOreproductive(age(women?(

(

Ilona!Nenko1,!Andrzej!Galbarczyk1,!Magdalena!Klimek1,!Pedro!Fernandez2,!Grazyna!Jasienska1!1Department!of!Environmental!Health,!Jagiellonian!University!Medical!College,!Krakow,!Poland!2Boston!University!School!of!Medicine,!Boston,!USA(!Objective:!Alzheimer’s!disease!is!the!most!common!type!of!dementia.!It!is!predicted!that!more!than!15!million!people!will!be!living!with!dementia!by!2050!and!higher!prevalence!is!expected!in!low!income!countries.!Therefore,!it!is!important!to!detect!as!many!risk!factors!for!dementia!as!possible.!In!women,!reproductive!


! 20!

processes!are!related!to!many!metabolic!changes,!including!increased!oxidative!stress.!Oxidative!stress!is!involved!in!the!onset!of!several!neurodegenerative!disorders.!We!hypothesize!that!high!lifetime!reproductive!investment!may!be!negatively!related!to!cognitive!functioning!in!older!women.!!Methods:!Our!study!examined!relationships!between!reproductive!history!traits!and!cognitive!functioning!assessed!by!MiniLMental!State!Examination!(MMSE)!and!clockLdrawing!test.!There!are!simple!test!of!cognitive!function!based!on!a!possible!score!of!30!points!(MMSE)!and!score!of!10!points!(clockLdrawing!test).!Scores!from!MMSE!were!standardised!for!age!and!years!of!education.!We!examined!378!rural,!parous!women!aged!45!to!92!years!(mean!=!61.4)!from!rural!population!with!a!high!birth!rate!from!Southern!Poland.!!Results:!Preliminary!analyses!indicate!that!number!of!children!born,!age!at!first!reproduction!and!age!at!last!reproduction!are!not!related!to!a!higher!risk!of!mild!cognitive!decline!in!women!(standardised!MMSE!score!below!29!points)!or!low!score!in!clockLdrawing!test.!Conclusion:!Reproductive!investment!is!not!related!to!cognitive!functioning!in!postLreproductive!age!women.!It!is!likely!that!support!provided!by!children,!especially!in!older!age,!counteracts!costs!of!reproduction!in!women.!!!!Environmental(impact(on(lifeOhistoryOstrategies(in(rotifers!!Heike!Gruber,!Lutz!Becks!MaxLPlanckLInstitute!for!Evolutionary!Biology,!Plön,!Germany!!Organisms!across!the!tree!of!life!pursue!a!wide!variety!of!lifeLhistoryLstrategies.!Differences!in!strategies!can!be!observed!by!monitoring!fitness!(lifespan!and!reproduction)!also!within!species.!We!investigate!whether!and!how!lifeLhistoryLstrategies!change!under!various!environmental!influences!in!a!freshLwater!rotifer.!The!goal!of!our!study!was!to!find!out!i)!which!lifeLhistoryLstrategies!are!followed!under!which!environmental!conditions!and!ii)!what!are!the!epigenetic!mechanisms!determining!the!strategies.!Therefore!different!clonal!populations!of!the!parthenogenetic!rotifer!Brachionus%calyciflorus!were!provided!with!different!energy!supply!levels!and!fitness!was!documented!for!three!generations.!We!found!significant!differences!in!life!history!traits!across!different!clonal!lines!and!depending!on!the!supplied!energy!level!and!thus!standing!genetic!variation!for!different!lifeLhistoryLstrategies!within!populations.!Furthermore,!we!observed!a!positive!correlation!of!energy!supply!rate!with!reproductive!output,!but!no!tradeLoff!between!measured!fitness!traits.!The!results!of!this!and!further!studies!with!our!novel!model!system!may!give!insights!into!the!underlying!mechanisms!for!the!different!lifeLhistoryLstrategies!and!reveal!the!impact!of!environmental!influences!on!strategies!also!in!other!organisms,!including!human.!!!How(cognition(and(sociality(have(shaped(human(biology:(a(biodemographic(perspective.(

Samuel!Pavard1!1!EcoHAnthropologie%et%Ethnobiologie,%UMR%7206%CNRS,%MNHN,%Univ%Paris%Diderot,%Sorbonne%Paris%Cité,%FH

75016%Paris,%France!!!!

The!use!of!tools!was!thought!to!have!emerged!less!than!2!MYA,!granting!their!makers!to!be!named!as!the!first!of!their!kind:!Homo%habilis.!However,!in!the!past!5!years,!new!archeological!evidence!has!shown!that!the!use!of!tools!has!appeared!long!before!3!MYA!with!Australopithecus%africanus.!Parallel!to!this,!genetic!and!archeological!data!finally!agree!to!date!humans–chimpanzees!divergence!around!6!MYA.!Consequently!hominins!have!spent!more!than!half!the!time!since!this!divergence,!not!discovering,!but!enhancing!tools!technology.!This!has!drastically!changes!our!view!on!human!evolution!because!this!time!was!enough!for!our!biology!to!evolve!in!response!to!the!use!of!tools.!For!example,!it!is!more!and!more!accepted!that!the!use!of!tools!has!led!to!a!change!in!diet!which!in!turn!had!deeply!impacted!brain!ontogeny!(Hublin!et!al.!2015).!!

Here!we!furthermore!argue!that!most!aspects!of!human!lifeLhistory!have!been!shaped!by!the!coevolution!between!biology,!cognition!and!sociality!in!our!species.!A!biodemographic!perspective!is!therefore!crucial!to!review,!organize!and!interpret!recent!theoretical!and!empirical!knowledge!in!various!fields,!from!paleoanthropology!to!neurology.!Together!this!sheds!new!light!on!crucial!human!evolutionary!questions!such!as!the!obstetric!dilemma!and!the!evolution!of!child!altriciality,!the!emergence!of!the!hazardous!but!yet!particularly!efficient!female!reproduction,!and!the!importance!of!kinship!for!transfers!of!resources!and!ideas.!(


! 21!

Evolutionary(demography(

(

Hans!Metz!Mathematical!Institute,!University!of!Leiden!

!The!mechanisms!of!population!change!reside!at!least!partially!in!the!demographic!behaviour!of!individuals.!

To!lift!mechanisms!at!the!individual!level!to!the!population!level!we!need!to!find!a!state!space!model!capturing!this!behaviour!and!a!representation!of!the!environment!affecting!the!state!transitions!and!the!behavioural!outputs!such!that!once!the!environment!is!given!this!effectively!decouples!individuals.!If!we!can!do!so!it!follows!that,!if!it!does!not!go!extinct!shortly!after!startLup!due!to!its!small!initial!size,!a!population!in!an!ergodic!environment!will!in!the!long!run!on!average!grow!or!decline!exponentially.!!!

More!often!than!not!individuals!affect!the!dynamics!of!the!environment!through!a!form!of!mass!action,!that!is,!through!the!added!contributions!of!individuals.!In!that!case,!if!sufficiently!many!individuals!are!exposed!to!a!similar!environment!we!can!approximate!the!population!dynamics!by!a!deterministic!dynamical!system.!I!shall!restrict!myself!to!such!populations!as!small!populations!do!not!persist!sufficiently!long!for!gradual!evolution!to!unfold.!!

In!the!real!world!populations!are!necessarily!bounded.!Hence,!communities!have!attractors,!which!usually!generate!ergodic!environments.!In!that!case,!clonally!reproducing!residents!necessarily!have!zero!long!term!growth!rate.!This!rule!extends!to!alleles!in!Mendelian!populations.!

Evolution!happens!thanks!to!the!fact!that!individuals!reproduce!almost!faithfully.!In!order!to!get!a!grip!on!the!resulting!changes!in!population!composition!I!assume!that!mutations!are!sufficiently!rare!that!they!do!not!interfere!during!their!substitution.!!

If!we!can!characterize!hereditary!types!with!continuous!traits!and!if!a!mutant's!demographic!behaviour!depends!smoothly!on!its!trait!vector,!then!the!asymptotic!per!capita!growth!rate!of!a!mutant!type!in!a!given!resident!environment!(its!"invasion!fitness")!depends!smoothly!on!the!mutant's!traits.!For!"good"!community!attractors!the!same!holds!good!for!the!dependence!on!the!traits!of!the!resident!types,!at!least!away!from!community!dynamical!bifurcations.!!

I!shall!for!now!assume!away!the!latter!complication.!This!sets!the!geometrical!structure!for!adaptive!dynamics,!a!dynamics!(AD)!generated!by!repeated!mutant!invasions!in!the!union!over!n!of!the!trait!space!to!the!power!n!(the!population's!evolutionary!state),!where!n!stands!for!the!current!degree!of!polymorphism.!!If!mutational!steps!are!small,!mutants!with!positive!invasion!fitness!substitute,!except!close!to!points!where!the!selection!gradient!(the!differential!of!the!invasion!fitness!for!the!mutant!trait)!equals!zero,!the!soLcalled!evolutionarily!singular!points.!This!property!allows!writing!down!for!each!n!a!differential!equation!for!the!trait!vectors!(the!soLcalled!Canonical!Equation!of!AD),!on!the!assumption!of!small!mutational!steps!and!away!from!those!singular!points!(which!for!a!given!n!also!are!the!stationary!points!of!the!CE).!!

The!latter!comprise!i.a.!the!attracting!ESSes!and!points!where!the!evolutionary!trajectory!branches.!

Sensitivity(of(population(growth(rate(to(changes(in(vital(rates:(A(comparison(of(transient(and(asymptotic(

dynamics(in(stationary(and(nonOstationary(environments(

Nora%Sánchez%Gassen%and%Hal%Caswell%

Institute%for%Biodiversity%and%Ecosystem%Dynamics,%University%of%Amsterdam%

In!an!article!from!1982,!Lande!showed!that!the!mean!fitness!of!a!population!can!be!equated!with!its!growth!rate.!When!analyzing!the!links!between!changes!in!the!vital!rates!and!population!growth!or!fitness,!evolutionary!biologists!have!so!far!mostly!focused!on!stable!populations!in!constant!environments,!or!stochastically!stable!populations!in!stationary!stochastic!environments,!to!understand!how!changes!in!a!vital!rate!affect!asymptotic!growth!rates.!Nonetheless,!natural!populations!are!rarely!stable!and!their!environments!not!always!stationary.!Environmental!catastrophes!or!climate!changes!may!produce!nonstationary!trends!in!a!population’s!vital!rates!and!lead!to!unstable!age!or!stage!distributions!and!fluctuating!growth!rates.!It!is!so!far!less!well!understood!how!changes!in!ageL!or!stageLspecific!vital!rates!influence!the!shortLterm!growth!of!transient!populations!in!nonstationary!environments.!

Here,!we!contribute!to!this!discussion!by!focusing!on!human!populations!in!Europe.!Populations!across!European!countries!have!undergone!substantial!changes!in!their!demographic!behavior.!During!the!last!fifty!years!alone,!fertility!rates!have!declined!across!Europe,!and!increased!again!more!recently!in!some;!mortality!


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rates!have!declined;!and!migration!numbers!have!fluctuated!substantially,!often!from!one!year!to!the!next.!These!changes!in!demographic!behaviour!are!ongoing,!and!are!expected!continue!in!the!future.!Sensitivity!analysis!can!help!us!identify!the!relationship!between!these!changes!in!the!vital!rates!and!transient!population!sizes!in!the!short!term.!We!use!a!set!of!countryLspecific!population!projections!recently!published!by!the!European!Statistical!Office!(Eurostat)!and!examine!the!sensitivity!of!projected!population!sizes!in!2080!to!perturbations!in!the!vital!rates,!applied!equally!in!each!projection!year.!The!results!reveal!to!what!extent!individual!vital!rates!influence!population!size!and!shortLterm!growth,!and!which!perturbations!would!lead!to!higher!final!population!sizes!and,!thereby,!increase!‘fitness’.!By!comparing!results!across!countries!we!can!also!identify!similarities!and!differences!in!transient!dynamics!across!populations.!!Mother’s(curse(and(the(evolution(of(sexOspecific(patterns(of(senescence(in(the(wild(

!Lemaître(JF,!Gaillard!JM,!Conde!T,!Tidière!M,!Lartillot!N!&!Marais!G.!%

Université%de%Lyon,%FH69000,%Lyon;%Université%Lyon%1;%CNRS,%UMR5558,%Laboratoire%de%Biométrie%et%Biologie%

Évolutive,%FH69622,%Villeurbanne,%France.!!

In!most!animal!species,!males!and!females!differ!markedly!in!their!life!expectancy!and!senescence!patterns.!For!instance!in!mammals,!females!show!a!much!longer!longevity!than!males.!In!line!with!this!observation,!a!stronger!rate!of!actuarial!senescence!is!often!described!in!mammalian!males.!Understanding!the!evolutionary!origins!of!these!sex!differences!in!actuarial!senescence!patterns!is!currently!an!important!challenge!in!ageing!research.!!

Several!hypotheses!have!been!proposed!to!explain!the!evolution!of!sexLspecific!senescence!patterns!(e.g.!heterogametic!sex!hypothesis,!sexual!selection)!but!here!we!will!focus!on!the!rarely!tested!‘Mother’s%curse%

hypothesis’.!In!eukaryotes,!mitochondrial!DNA!is!inherited!from!the!mother.!Therefore,!mitochondrial!mutations!can!be!accumulated!in!the!genome!even!if!they!have!maleLspecific!deleterious!effects.!For!instance,!this!has!been!shown!in!humans!where!some!mitochondrial!mutations!are!associated!with!male!infertility.!!

Here,!we!have!tested!the!hypothesis!that!a!higher!rate!of!mutations!in!mitochondrial!DNA!is!responsible!for!an!earlier!and/or!stronger!senescence!in!males!compared!to!females.!To!address!this!question,!we!used!a!dataset!of!about!100!mammalian!species!for!which!we!gathered!both!genomic!data!(e.g.!mitochondrial!DNA!mutation!rate)!and!sexLspecific!demographic!data!(ageLspecific!mortality!estimates!for!both!males!and!females).!Overall,!our!comparative!study!provides!new!insights!into!the!factors!shaping!sex!differences!in!actuarial!senescence!patterns!across!the!tree!of!life.!!Risk(Perception’s(Influences(on(Teenage(Childbearing(–(A(Genetic(Perspective(

Lin!Chen!!Department!of!Sociology,!University!of!Oxford(!

This!research!seeks!to!answer!the!following!research!question:!In!a!geneLenvironment!interaction!perspective,!is!there!any!genetic!correlation!between!risk!perception!and!teenage!childbearing?!This!paper!is!going!to!utilize!candidate!geneLenvironment!analysis!to!distinguish!effect!of!genetic!baseline!risk!perception!on!teenage!childbearing!from!environmental!effect!and!geneLenvironment!interaction!on!teenage!childbearing.!Cox!model!with!a!piecewise!constant!intensity!is!applied!to!analysis!the!geneLenvironment!interaction!of!risk!perception’s!genetic!influence!on!different!age!groups!of!teenagers!(<=15,!16L17,!18L19).!This!research!is!based!on!data!set!from!The!National!Longitudinal!Study!of!Adolescent!Health!(Add!Health)!with!Candidate!Genes!Data!on!risk!perception.!By!answering!the!research!question!and!further!untangling!the!underline!relation!between!risk!perception!and!teenage!childbearing,!the!value!of!this!research!has!twoLfold.!Theoretically,!this!research!contributes!a!genetic!perspective!on!the!influence!of!risk!perception!on!teenage!childbearing!perception!by!employing!the!genetic!component.!Methodologically,!by!applying!the!geneLenvironment!analysis,!this!research!unfolds!the!mediating!mechanism!between!risk!perception!and!teenage!childbearing!into!three!dimensions:!genetic,!environment!and!geneLenvironment!interaction!thereby!filling!the!gap!of!previous!research,!which!mainly!focuses!on!the!social!environment!effect.!!!!!


! 23!

Wind(is(an(important(climatic(factor(

!Tina!Cornioley1,!Luca!Börger2,!Arpat!Ozgul1,!Henri!Weimerskirch3!1!Institute!of!Evolutionary!Biology!and!Environmental!Studies,!University!of!Zurich,!Switzerland!2!Biosciences,!Swansea!University,!UK!3!Centre!d'Études!Biologiques!de!Chizé,!Centre!National!de!la!Recherche!Scientifique,!France!!

Wind!is!an!important!climatic!factor!for!flying!animals!as!by!affecting!their!locomotion!it!can!deeply!impact!their!life!history!characteristics.!Species!responses!to!wind!speed!however!have!not!received!much!attention!although!the!global!wind!patterns!are!changing.!

We!built!a!foraging!model!describing!the!relationship!between!the!wind!and!foraging!pattern!of!the!wandering!albatross,!a!pelagic!seabird!which!mass!has!increased!in!parallel!to!wind!speed!increase!over!the!past!decades.!We!focused!on!incubation!period,!the!most!critical!reproductive!stage!for!this!species.!

By!linking!wind!and!foraging!outcome,!the!foraging!model!allowed!predicting!breeders'!mass!over!time!at!different!wind!speed.!We!simulated!the!body!mass!of!incubating!pairs!and!their!incubation!success!under!varying!wind!speed!scenarios.!Incubation!failure!is!usually!due!to!nest!abandonment!triggered!by!substantial!mass!loss.!We!tracked!the!number!of!times!critical!mass!leading!to!nest!abandonment!was!reached!to!assess!incubation!success.!

As!expected!for!a!species!performing!dynamic!soaring,!wind!speed!increased!bird!ground!speed.!However,!the!simulation!predicted!no!overall!effect!of!wind!speed!on!mass!over!the!incubation!period!of!wandering!albatrosses!and!no!nest!abandonment.!

We!found!indications!that!wandering!albatrosses!behave!as!time!minimizer!during!incubation!as!mass!gain,!a!proxy!for!energy!intake,!was!independent!of!trip!duration!and!ground!speed.!Time!minimizers,!in!contrast!to!energy!maximizes!which!forage!for!a!fixed!amount!of!time,!forage!until!their!energy!requirements!are!fulfilled.!Furthermore,!lighter!individuals!gained!more!mass,!indicating!that!wandering!albatrosses!adjust!energy!requirement!to!their!body!conditions.!This!can!come!at!the!cost!of!their!partner's!conditions!as!mass!loss!of!the!incubating!partner!depended!on!trip!duration.!This!behaviour!is!consistent!with!strategies!of!longLlived!species!which!favoured!their!own!survival!over!their!current!reproductive!success.!!!Evaluating(the(fitness(importance(of(maternal(size(effects(on(offspring(in(fish:(An(integral(projection(model(

(

Yngvild!Vindenes§*,!Øystein!Langangen§,!Ian!J.!Winfield$,!Asbjørn!Vøllestad§%%

§Centre!for!Ecological!and!Evolutionary!Synthesis!(CEES),!Department!of!Biosciences,!University!of!Oslo,!Norway!$Lake!Ecosystems!Group,!Centre!for!Ecology!&!Hydrology,!Lancaster,!U.K.!*[email protected]!!

Demographic!rates!of!fish!often!depend!on!body!size,!including!parameters!affecting!reproduction!such!as!fecundity,!egg!size,!and!probability!of!maturity.!Recent!studies!have!reemphasized!the!potential!importance!of!maternal!body!size!to!early!survival!and!growth!of!offspring,!through!various!mechanisms.!!For!instance,!large!female!fish!tend!to!produce!larger!eggs,!assumed!to!result!in!more!”high!quality”!offspring!(i.e.,!with!higher!survival!and/or!increased!growth!rates).!This!suggests!that!fishery!management!may!need!to!account!for!the!population!size!structure!in!addition!to!biomass,!and!apply!size!limits!that!preserve!the!largest!females!as!well!as!the!small.!!

We!tested!the!importance!of!such!maternal!size!effects!to!fitness!using!a!femaleLbased!integral!projection!model!(IPM)!for!a!population!of!pike!(Esox%lucius)!from!Windermere,!UK,!where!unique!longLterm!data!are!available.!The!model!includes!length,!length!at!age!1,!and!temperature!as!state!variables,!and!accounts!for!individual!heterogeneity!in!growth!and!survival!mediated!by!length!at!age!1.!Average!egg!weight!increased!with!maternal!length,!leveling!off!for!the!largest!lengths.!However,!a!sensitivity!analysis!revealed!that!this!maternal!effect!had!only!minor!impacts!on!the!growth!rate!lambda!(fitness),!even!if!we!assumed!a!very!strong!positive!effect!of!egg!weight!on!offspring!survival.!In!contrast,!lambda!was!sensitive!to!temperature!conditions!during!the!first!year,!having!longLlasting!fitness!consequences!through!growth.!!

The!results!do!not!support!the!hypothesis!that!large!females!contribute!more!to!population!growth!(relative!to!their!biomass),!but!instead!highlight!the!importance!of!environmental!conditions!experienced!in!


! 24!

early!life.!!We!believe!these!results!can!be!generalized!to!species!of!similar!life!histories,!including!many!commercially!important!fishery!species.!The!model!provides!useful!knowledge!to!conservation!and!management!of!such!species,!including!actions!to!mitigate!impacts!of!climate!warming.!!!!

What(are(the(consequences(of(early(life(stress(for(fertility(in(human(populations?((

Rebecca!Sear1,!Cristina!Moya2!&!Paula!Sheppard1!1!London!School!of!Hygiene!&!Tropical!Medicine,!UK!2!University!of!British!Columbia,!Canada!

There!has!been!a!recent!flurry!of!literature!in!the!human!evolutionary!sciences!suggesting!that!early!life!stress!can!kickstart!a!‘fast’!life!history,!including!earlier!age!at!first!reproduction!and!higher!total!numbers!of!children!born.!Most!evidence!for!this!hypothesis!comes!from!data!correlating!early!life!stress!with!earlier!first!births!in!women.!Other!measures!of!‘life!history!strategy’!are!rarely!investigated.!This!talk!will!consider!how!the!wide!availability!of!data!from!a!range!of!human!populations!can!be!harnessed!to!answer!the!question!of!whether!early!stress!does!‘speed!up’!life!history!strategy.!Particular!emphasis!will!be!placed!on!the!loss!of!a!parent(s)!as!an!indicator!of!stress,!and!measures!of!fertility!as!indicators!of!life!history!strategy.!Results!suggest!that!in!high!income,!low!fertility!populations,!early!life!stress!is!consistently!associated!with!an!early!age!at!first!birth!for!women,!but!may!have!little!impact!on!other!measures!of!fertility!such!as!total!number!of!children!born.!Associations!between!early!life!stress!and!subsequent!reproductive!outcomes!are!less!consistent!in!low!income,!high!fertility!populations,!though!fewer!studies!have!been!conducted!in!such!populations.!We!conclude!that!the!question!of!whether!early!life!stress!is!correlated!with!a!‘faster’!life!history!strategy,!including!early!first!births!and!higher!overall!fertility,!is!still!open.!Further!empirical!investigation!is!required!to!both!explore!these!relationships!in!a!wider!range!of!ecological!contexts,!and!to!determine!the!causal!mechanisms!by!which!stress!may!influence!fertility!and!other!life!history!outcomes.!!!!The(evolution(of(maternal(effect(senescence(

Jacob!Moorad!&!Daniel!Nussey!Institute!of!Evolutionary!Biology,!University!of!Edinburgh,!Charlotte!Auerbach!Road,!Edinburgh!EH9!3FL!!

Increased!maternal!age!at!reproduction!is!often!associated!with!decreased!offspring!performance!in!numerous!species!of!plants!and!animals!(including!humans).!Current!evolutionary!theory!considers!such!maternal!effect!senescence!as!part!of!a!unified!process!of!reproductive!senescence,!which!is!under!identical!ageLspecific!selective!pressures!to!fertility.!We!offer!a!novel!theoretical!perspective!by!combining!William!Hamilton’s!evolutionary!model!for!aging!with!a!quantitative!genetic!model!of!indirect!genetic!effects.!We!demonstrate!that!fertility!and!maternal!effects!senescence!are!likely!to!experience!different!patterns!of!ageLspecific!selection!and!thus!can!evolve!to!take!divergent!forms.!Applied!to!neonatal!survival,!we!find!that!selection!for!maternal!effects!is!the!product!of!ageLspecific!fertility!and!Hamilton’s!ageLspecific!force!of!selection!for!fertility.!Population!genetic!models!show!that!senescence!for!these!maternal!effects!can!evolve!in!the!absence!of!reproductive!or!actuarial!senescence;!this!implies!that!maternal!effect!aging!is!a!fundamentally!distinct!demographic!manifestation!of!the!evolution!of!aging.!However,!brief!periods!of!increasing!maternal!effects!can!evolve!when!fertility!increases!with!age!faster!than!cumulative!survival!declines.!This!is!most!likely!to!occur!early!in!life.!Our!integration!of!theory!provides!a!general!framework!with!which!to!model,!measure,!and!compare!the!evolutionary!determinants!of!the!social!manifestations!of!aging.!Extension!of!our!maternal!effects!model!to!other!ecological!and!social!contexts!could!provide!important!insights!into!the!drivers!of!the!astonishing!diversity!of!lifespans!and!aging!patterns!observed!among!species.!!!DemographyObased&assessment&of&bird&population&trends!

Sharon'Janssen,'Eelke%Jongejans!Radboud'University,'Nijmegen,'NL!

Many%bird%species%need%protection%because%their%population%sizes%are%declining.%The%priorization!of#which#bird%species%need%protection%most%is%mostly%based%on%the%categorization%made%with%the%Red%List%system.%With%this%method%species,%which%have%a%significant%chance%of%decline%in%the%future%but%which%are%not%declining%yet,%


! 25!

are$not$likely$to$be$protected.%For%such%reasons%there%is%need%for%a%more%scienceLbased&framework&for&bird&protection.*Here*we*evaluate*the*use*of*stochastic*population*models*for*analyzing*current*trends*in*national*population)data)of)bird)species)in)the)Netherlands.)The)case)studies#of#the#chiffchaff#and#wren#show#a#promising)role)for)stochastic)population)models)for)interpreting)current)trends)in)national)bird)data.)Based)on)parameterized*population*models*we*show*which*observed*declines*are*alarming*and*which*can*be*expected*for$a"fluctuating"but"longLterm%stable%population.%We%discuss%several%issues%with%these%analyses%and%prospects%for$traitLbased&interpolations&for&other&bird&species.!!Environmental(drivers(of(metapopulation(dynamics(in(the(spatially(and(socially(structured(Siberian(jay(

(

Kate!LaytonLMatthews,!!Arpat!Ozgul!Universität!Zürich,!Zürich,!Switzerland!!

How!a!population!responds!to!environmental!change!will!determine!its!persistence.!By!looking!at!lifeLhistory!traits!in!a!population!of!socially!structured!Siberian!jays,!Perisoreus%infaustus,!we!investigate!how!susceptible!the!population!is!to!environmental!variability.!Siberian!jays!live!in!small,!social!groups,!within!relatively!stable!territories.!The!study!site!in!Northern!Sweden!offers!an!excellent!opportunity!to!study!the!relationship!between!demography!and!environment!due!to!its!seasonal!climate!and!spatially!heterogeneous!landscape!(due!to!commercial!forestry).!Previous!research!has!already!shown!the!negative!impact!of!forestry!on!specific!lifeLhistory!components,!but!its!influence!on!the!population!as!a!whole!is!as!of!yet!unexplored.!

We!modeled!demographic!rates!(survival,!maturation,!breeding!probability!and!reproductive!success)!using!25Lyears!of!markL!recapture!data!from!1344!individuals.!The!environmental!covariates!include!both!temporal!measures!that!describe!the!climate!and!changes!in!habitat!use!as!well!as!spatial!measures!of!habitat!quality.!The!measures!of!habitat!quality!were!chosen!based!on!those!considered!most!representative!of!the!changes!resulting!from!forest!management.!!

Our!analysis!so!far!has!shown!that!seasonality,!climatological!factors!and!densityLdependence!explain!considerable!temporal!variation!in!stageLspecific!survival!and!reproductive!success.!Although!both!survival!and!reproductive!success!appear!to!deteriorate!in!habitats!undergoing!forestry,!no!measures!of!habitat!quality!elucidated!the!cause!of!this!decline.!One!potential!explanation,!increased!predation!risk,!is!currently!under!investigation.!The!final!step!is!to!develop!a!stageLstructured,!matrixLbased!metapopulation!model!to!investigate!how!the!population!responds!to!the!observed!changes!in!demographic!rates!and!how!the!underlying!environmental!processes!are!driving!population!dynamics.!This!spatiallyL!explicit!model!will!link!local!population!dynamics!that!vary!based!on!habitat!quality!with!stageLspecific!dispersal!!Introducing(hyperOmatrix(models(

Gregory!Roth!and!Hal!Caswell(Institute%for%Biodiversity%and%Ecosystem%Dynamics,%University%of%Amsterdam,%1090GE%Amsterdam,%Netherlands%

!Demographic!models!describe!population!dynamics!in!terms!of!the!states!of!the!individuals!making!up!the!

population!(size,!age,!developmental!stage,!parity,!frailty,!physiological!condition);!these!are!called!iHstates!(Metz!and!Diekmann!1986).!The!model!itself!is!formulated!in!term!of!the!movement!of!individuals!among!iLstates.!

Classical!demographic!models!classified!individuals!by!a!single!criterion.!This!was!enlarged!to!two!criteria!in!vecLpermutation!models!and!the!closely!related!megamatrix!models.!However,!it!has!been!recognised!that!the!interplay!of!more!than!two!criteria!can!affect!the!population!dynamics;!for!example,!in!models!for!migration!including!age,!size,!and!spatial!location;!in!models!for!reproductive!performance!including!age,!parity,!and!duration;!in!models!for!mortality!including!age,!stage,!and!frailty;!in!stochastic!models!including!size,!frailty,!and!environmental!state.!

Here,!we!present!a!model!framework!in!which!individuals!may!be!classified!by!any!number!of!criteria,!using!the!generalisation!of!the!vecLpermutation!approach!to!hyperLmatrices.!These!models!are!constructed!from!a!simple!blockLdiagonal!matrix!formulation!of!the!movement!of!individuals!among!each!dimension!of!the!iLstate.!This!formulation!makes!available!the!usual!demographic!analysis!developed!for!classical!matrix!models.!The!possible!applications!include!the!models!mentioned!above!but!are!not!limited!to!three!criteria.!

Sensitivity!analysis!calculates!the!effect!of!parameters!changes!on!any!demographic!outcomes!of!a!model!(Caswell!2001).!Demography!and!evolution!meet!when!the!outcome!of!interest!is!the!fitness!of!a!phenotype;!


! 26!

its!sensitivity!to!any!vector!of!traits!provides!the!selection!gradient!on!those!traits.!Here,!we!derive!a!general!formula!for!the!sensitivity!of!any!output!of!the!hyperLmatrix!model,!to!any!vector!of!parameters.!

!MegaOanalysis(of(31,396(individuals(beyond(reproductive(age(unravels(strong(GxE(interaction(on(human(

fertility(

Felix!C!Tropf,!Renske!M!Verweij,!Peter!J!van!der!Most,!Gert!Stulp,!Maja!Djundeva,!Anastasia!Nyman,!Sarah!Medland,!Nick!Martin,!Peter!M!Visscher,!Harold!Snieder,!S!Hong!Lee,!Melinda!C!Mills!!

A!megaLanalysis!applying!whole!genome!methods!and!genetic!profile!score!analysis!on!31,396!men!and!women!from!seven!Western!populations!unravels!strong!gene!–!environment!interaction!on!human!fertility.!Heritability!estimates!based!on!molecular!genetic!data!of!the!whole!genome!suggested!that!only!4!per!cent!(SE!1%)!of!the!variance!in!number!of!children!ever!born!(NEB)!and!5!per!cent!(SE!1%)!for!AFB!are!explained!by!genetic!difference!amongst!unrelated!individuals.!The!gene!–!environment!interaction!models,!however,!show!that!for!individuals!from!the!same!population!who!live!in!the!same!time!period!before!or!after!the!Second!World!War,!the!estimated!genetic!variance!is!(more!then)!four!fold!higher!with!23!per!cent!(SE!3%)!for!NEB!and!19!per!cent!(SE!4%)!for!AFB.!Genetic!profile!scores!unravel!that!genes!associated!with!human!height!have!different!effects!on!the!NEB!across!populations!and!genes!associated!with!educational!attainment!have!approximately!doubled!effects!on!women’s!AFB!across!the!past!century.!Our!findings!suggest!first!that!evolutionary!forecasts!due!to!natural!selection!on!human!populations!are!probably!not!possible!due!to!rapidly!changing!selection!pressure,!second!that!gene!environment!interaction!is!important!to!explain!low!heritability!findings!in!fertility!beyond!family!studies!and!third!that!psychological!or!cognitive!traits!potentially!mediate!genetic!effects!on!fertility.!Our!findings!call!for!interdisciplinary!work!to!understand!realized!human!fertility.!!!!Climate,(rather(than(harvest,(is(the(main(driver(of(ageOspecific(mortality(trajectories(in(a(tropical(tree(

!Orou!G.!Gaoue1,!Carol!C.!Horvitz2!and!Shripad!Tuljapurkar3!1!Department!of!Botany,!University!of!Hawaii!at!Manoa,!Honolulu,!HI!96822,!USA!2!Department!of!Biology,!University!of!Miami,!Coral!Gables,!FL!33146,!USA!3!Department!of!Biology,!Stanford!University,!Stanford,!CA!94305,!USA.!!

A!mechanistic!understanding!how!environmental!and!anthropogenic!factors!interact!(e.g.,!drought,!harvest!intensity!or!herbivory)!to!shape!population!demography!is!central!to!designing!resource!exploitation!and!management!plans.!For!longLlived!plants,!theoretical!and!empirical!studies!demonstrate!that!mortality!represents!the!most!sensitive!and!important!demographic!process!that!determine!the!ability!of!a!species!to!persist!in!an!environment!and!respond!to!anthropogenic!pressure.!!We!used!ageLfromLstage!matrix!modeling!to!investigate!how!severe!foliage!(for!fodder)!and!bark!(for!medicine)!harvest!by!indigenous!people!in!two!climatically!different!regions!(dry!versus!moist)!affects!ageLspecific!mortality!trajectories!of!a!tropical!tree,!Khaya%senegalensis!in!West!Africa.!We!compared!models!with!constant!harvest!intensity!over!time!and!those!with!stochastically!variable!harvest!intensity.!The!effect!of!harvest!on!ageLspecific!mortality!trajectories!was!weak!and!only!noticeable!in!the!moist!region.!However,!ageLspecific!mortality!trajectories!differed!between!regions.!In!the!moist!region,!mortality!rates!decreased!with!age!for!the!first!30!years!to!a!minimum!rate!and!then!increased!gradually!after!to!reach!the!mortality!plateau.!In!the!dry!region,!mortality!rates!decreased!with!age!to!reached!the!plateau!asymptotically.!This!difference!in!ageLspecific!mortality!trajectory!is!due!to!a!greater!delay!to!reaching!reproductive!size/age!in!the!dry!region.!Our!findings!underscore!intraspecific!variation!in!ageLspecific!mortality!schedules!and!indicate!that!climatic!effect!may!override!the!impact!of!anthropogenic!activities!on!plant!demography.!eLmail:[email protected]!! !


! 27!

!!Consequences(of(pollen(limitation(for(plant(population(dynamics.(

(

Tiffany!Knight!Helmholtz!Centre!for!Environmental!Research,!Martin!Luther!University!HalleLWittenberg!Department!of!Biology,!Washington!University,!St.!Louis,[email protected]!!Wild!pollinators!have!declined!in!abundance!and!diversity,!and!there!is!global!concern!that!this!might!result!in!lower!reproductive!success!and!lower!rates!of!population!growth!for!wild!and!cultivated!plant!species.!!We!have!conducted!a!metaLanalysis!of!pollen!supplementation!experiments!in!wild!plant!populations!and!find!that!pollen!limitation!is!common!and!that!the!magnitude!of!pollen!limitation!can!be!quite!high.!!Here,!we!consider!the!relationship!between!pollen!limitation!and!plant!population!growth!rate!and!discuss!the!conditions!for!which!pollen!limitation!is!a!conservation!concern.!!Pollen!limitation!is!most!likely!to!cause!a!decline!in!plant!population!growth!rate!when!the!magnitude!of!pollen!limitation!is!high,!when!population!dynamics!are!sensitive!to!changes!in!fecundity,!when!current!reproductive!output!has!little!effect!on!survivorship,!growth,!and/or!future!reproduction,!and!when!populations!are!seed!limited.!!We!conclude!that!information!on!the!life!history,!breeding!system!and!habitat!association!of!the!plant!species!can!be!used!as!a!guide!to!interpreting!the!consequences!of!pollen!limitation!for!plant!population!dynamics.!!

Wednesday,(7(October(2015(

! 28!

!(

! Presenter!! Title!(abridged)! !Breakfast!! Board!meeting,!Evolutionary!Demography!Society!09:10L09:20! Marjolijn!Das! The!attraction!of!the!city:!female!ruralLurban!migration...!09:20L09:30! James!Anderson! Telomeres!and!longevity!09:30L09:40! Matthias!Galipaud! Optimal!mate!choice!strategies!under!reproductive!senescence!09:40L10:00! Keynote%9:!Annette!

Baudisch!The!molding!of!evolutionary!theories!of!aging!

10:00L10:10! Bruce!Kendall! A!natural!history!of!rarity!may!predict!recovery!...!10:10L10:20! Eldar!Rakhimberdiev! Evolutionary!machinery!of!rapid!body!size!decline...!10:20L11:00!!

Coffee(break(

11:00L11:10! Takenori!Takada! Elasticity!analysis!in!random!matrices!11:10L11:20! Stefano!Giaimo! Equilibria!in!a!population!with!ageLspecific!behaviors!11:20L11:50(

Keynote%10:!Jim!Vaupel!Lifespan!equality!and!life!expectancy!in!humans!and!other!primates!

11:50L12:10! Closing!remarks!12:10L13:20!!

Lunch!(shuttle!service!to!Lunteren!train!station!from!12:30L13:15)!

13:20L14:30!!

Workshops(1(and(2(

14:50L14:10! Coffee(break(

15:00L16:20! Workshops(1(and(2((continued)(

(

End!of!the!workshops,!shuttles!for!Lunteren!train!station!between!16:15!and!17:15!(

(

(

The(attraction(of(the(city:(Female(ruralOurban(migration(in(the(Netherlands(

!!Marjolijn!Das!!&!Jan!Latten!(Statistics!Netherlands,!University!of!Amsterdam)!!!Objective:(Nowadays,!in!the!largest!Dutch!cities!young!women!outnumber!young!men.!Migration!to!cities!is!often!triggered!by!opportunities!for!education!and!work.!We!investigate!whether!female!ruralLurban!migration!also!serves!to!increase!the!probability!of!finding!a!highLstatus!partner.!Women!derive!fitness!benefits!from!choosing!partners!with!a!high!social!status!and!a!high!level!of!resources!(Geary,!2004),!and!highLstatus!men!are!found!predominantly!in!the!cities!(Edlund,!2005).!(Methods:(The!study!is!based!on!five!integral!birth!cohorts!of!Dutch!women,!born!between!1966L1971!!in!less!urbanised!and!rural!areas!(N=287621).!Outcomes!were!measured!at!age!40L45.!Data!were!derived!from!administrative!registers!available!at!Statistics!Netherlands.!Results:(Compared!to!women!who!lived!in!less!urbanised!or!rural!areas,!women!aged!40L45!who!had!migrated!to!large!cities!were!more!often!highly!educated!and!had!higher!incomes.!!Regardless!of!their!own!educational!level,!women!more!often!had!highly!educated!partners!when!they!had!migrated!to!cities.!Lower!educated!women!who!had!migrated!were!more!often!childless!than!women!who!had!stayed!behind!in!more!rural!areas.!!Conclusions:Female!migration!to!cities!could!be!a!investment!in!their!own!resources!(education,!career)!as!well!as!an!investment!in!the!search!for!a!high!status!partner.!In!terms!of!reproductive!success,!migration!to!cities!might!be!a!risky!strategy!for!some:!lower!educated!city!women!had!a!higher!probability!of!being!childless!at!age!40L45.!!!!!!!


! 29!

Telomeres(and(Longevity(

James%J.%Anderson%%

University!of!Washington!!

Recent!hypotheses!propose!human!longevity!is!influenced!by!the!length!of!telomeres!(1).!Telomere!length!(TL)!undergoes!progressive!attrition!in!replicating!human!somatic!cells!in!culture.!Short!TL!ultimately!triggers!replicative!senescence.!The!same!process!ostensibly!occurs!in!replicating!tissues!of!the!human!soma.!!Thus,!at!any!age!TL!might!explain!in!part!the!number!of!cell!divisions!remaining!to!replicative!senescence.!Long!TL!allows!more!cell!divisions!and!bolster!tissue!repair!and!putatively!provides!the!immune!system!with!greater!reserves!to!fight!infection.!The!risk!of!atherosclerosis,!a!major!human!degenerative!disease,!is!increased!with!short!TL,!while!the!risk!of!major!cancers!is!increased!with!long!TL.!!The!proposed!mechanism!for!increased!cancer!risk!is!that!premalignant!clones!reach!larger!size!in!individuals!with!longer!TL,!raising!the!probability!that!the!clone!receives!additional!mutations!that!promote!malignant!transformation!(2).!!It!follows!that!TL!might!mediate!tradeoff!between!competing!sources!of!mortality!that!occur!at!differing!ages!and!under!different!levels!of!stress!and!environmental!conditions.!The!talk!describes!a!rudimentary!model!for!this!tradeLoff!in!which!the!rates!of!infection!and!degenerate!disease!accelerate!with!age!as!TL!diminishes!while!the!rate!of!!mutational!diseases!decelerate!as!TL!attrition!limits!the!replicative!potential!of!premalignant!clones.!Heritable!differences!in!TL!might!represent!adaption!to!differing!strengths!of!degenerative!and!mutational!sources!of!mortality!in!demographic/racial!groups.!The!framework!has!elements!found!in!antagonistic!pleiotropy!and!mutation!accumulation!theories!of!natural!selection.!

1. Aviv!A,!Kark!JD,!&!Susser!E!(2015)!Telomeres,!Atherosclerosis,!and!Human!Longevity:!A!Causal!Hypothesis.!Epidemiology%(Cambridge,%Mass.)!26(3):295L299.!

2. 2.!Aviv!A,!Anderson!JJ,!&!.Shay!JW!(in!review)!Aging!and!the!CancerLTelomere!Length!Paradox!(eLife).!

!Optimal(mate(choice(strategies(under(reproductive(senescence(through(accumulation(of(damages(

(

Matthias!Galipaud!a,!JeanLMichel!Gaillard!b,!JeanLFrançois!Lemaître!b!!

aBielefeld!University,!Department!of!Evolutionary!Biology,!Morgenbreede!45,!33615!Bielefeld.!!bUniversité!Claude!Bernard!Lyon!1,!UMR!CNRS!5558!LBBE,!43!bd!du!11!novembre!1918!69622!Villeurbanne.!!

Accumulation!of!somatic!damages!over!life!can!lead!to!a!decrease!in!fecundity!at!advanced!age.!Considering!that!damages!result!from!costly!actions!taken!by!animals!earlier!in!life,!individuals!should!tradeLoff!their!early!investment!in!reproduction!with!damage!accumulation!and!its!consecutive!fecundity!loss!in!late!life.!We!study!optimal!investment!in!mate!choice!by!females!over!life!using!a!stateLdependent!matrix!population!model.!In!our!model,!higher!levels!of!choosiness!provide!females!with!access!to!higher!quality!males!and!thus!to!increased!offspring!survival.!However,!because!it!leads!to!the!accumulation!of!damages,!choosiness!is!also!costly!in!terms!of!decreased!fecundity!and!of!increased!mortality!later!in!life.!Females!investing!heavily!in!mate!choice!early!in!life!therefore!enjoy!high!offspring!survival!but!rapidly!produce!fewer!young!and!died!prematurely.!On!the!other!hand,!females!investing!less!in!choosiness!suffer!overall!high!offspring!mortality!but!accumulated!only!little!damages,!remained!fecund!longer!and!died!later.!When!high!quality!males!provide!similarly!high!survival!probability!to!every!offspring!produced!by!a!female!at!a!reproductive!event!(e.g.!good!genes),!we!predict!that!females!should!increase!choosiness!with!age!due!to!terminal!investment.!However!when!benefits!provided!by!males!are!equally!divided!among!offspring!(e.g.!paternal!care),!females!decrease!their!choosiness!with!age.!Females!indeed!produce!fewer!offspring!when!they!become!older!and!do!not!need!high!quality!males!anymore!to!ensure!that!each!offspring!survives.!We!discuss!the!consequences!of!these!predictions!in!terms!of!population!ageLassortative!mating!and!strength!of!sexual!selection.!!!!!!!!!! !


! 30!

!

The(molding(of(evolutionary(theories(of(aging(

Annette!Baudisch!Department!of!Biology!MaxLPlanck!Odense!Center!for!the!Biodemography!of!Aging!University!of!Southern!Denmark!baudisch@biologyu.sdu.dk!!

The!seminal!paper!on!An%unsolved%problem%in%Biology!by!Sir!Peter!Medawar!in!1952!can!be!viewed!as!the!birth!of!the!evolutionary!theories!of!aging.!Since!1952,!the!theory!has!developed,!grown,!and!matured:!evidence!has!been!gathered!to!the!pro!and!to!the!contra,!new!tools,!methods!and!data!have!been!acquired,!and!new!directions!and!extensions!have!been!suggested!and!explored.!Now,!at!the!age!of!63,!the!original!theories!are!approaching!retirement!age.!It!is!time!to!let!them!resign!from!active!working!life!and!instead!enjoy!their!offspring!and!grandoffspring.!!A(natural(history(of(rarity(may(predict(recovery(from(endangerment(

!Bruce!E.!Kendall!!Bren!School!of!Environmental!Science!&!Management,!University!of!California,!Santa!Barbara,!USA!!

Efficient!conservation!with!limited!budgets!requires!triage,!directing!resources!to!programs!that!generate!the!most!conservation!benefit.!An!important!question!is!thus:!“Which!species!are!most!likely!to!respond!to!conservation!intervention?”!Demographic!theory!allows!us!to!identify,!from!a!sufficiently!well!quantified!life!history,!the!life!stages!and!vital!rates!most!responsive!to!threatLreducing!actions.!I!propose!that!knowing!something!about!a!species’!evolutionary!history!may!also!provide!guidance,!even!in!the!absence!of!intensive!demographic!study.!While!a!species!cannot!be!selected!to!be!rare,!a!population!that!is!forced!to!be!rare!will!experience!selection!on!traits!that!enhance!persistence.!Models!relating!extinction!time!to!carrying!capacity!suggest!that!any!species!that!is!naturally!rare!(i.e.,!has!persisted!for!a!long!time!with!a!low!carrying!capacity)!is!likely!to!have!a!life!history!that!exhibits!rapid!population!growth!at!low!density,!low!sensitivity!to!various!sources!of!environmental!stochasticity,!or!both—a!rare!species!without!such!characteristics!would!likely!have!gone!extinct!long!before!humans!began!to!affect!it.!Such!species!might!be!expected!to!recover!quickly!if!the!anthropogenic!factors!that!now!hold!them!below!their!natural!carrying!capacity!can!be!removed.!The!rapid!recovery,!after!threat!abatement,!of!species!native!to!small!islands,!such!as!the!Lord!Howe!Island!woodhen!(Australia)!and!the!Channel!Islands!fox!(California,!USA),!is!highly!suggestive!in!this!regard.!Using!simple!demographic!models,!I!show!how!naturally!enforced!rarity!rapidly!prunes!the!parameter!space!to!values!that!enhance!persistence,!and!that!species!that!have!survived!a!period!of!rarity!are!more!likely!to!recover!from!very!low!abundance!than!are!species!that!have!not!experienced!rarity.!Thus,!prioritizing!endangered!species!with!a!natural!history!of!rarity!may!result!in!the!greatest!likelihood!of!conservation!success.!!Evolutionary(machinery(of(rapid(bodyOsize(decline(in(a(migratory(bird(

!Eldar!Rakhimberdiev1,2!&!Theunis!Piersma1,3!1.!Department!of!Marine!Ecology,!NIOZ!Royal!Netherlands!Institute!for!Sea!Research,!PO!Box!59,!1790!AB,!Den!Burg,!Texel,!The!Netherlands!2.!Department!of!Vertebrate!Zoology,!Biological!Faculty,!Lomonosov!Moscow!State!University,!119991,!Moscow,!Russia!3.!Conservation!Ecology!Group,!Groningen!Institute!for!Evolutionary!Life!Sciences!(GELIFES),University!of!Groningen,!P.O.!Box!11103,!9700!CC!Groningen,!The!Netherlands!

!Female!BarLtailed!godwits!Limosa%lapponica%lapponica!wintering!in!the!Wadden!Sea!and!breeding!in!Scandinavia!have!declined!in!body!size.!BodyLshrinkage!is!now!seen!as!a!common!response!to!climate!change,!especially!in!Arctic!animals.!With!15!years!of!captureLresight!data!we!estimated!nonlinear!bivariate!fitness!(survival)!surfaces!for!these!birds!and!showed!strong!directional!selection!for!shortening!bill!length.!Seasonal!survival!captureLrecapture!models!localized!this!selection!(mortality!biased!towards!the!longer!billed!females)!


! 31!

on!the!wintering!grounds!in!the!Wadden!Sea!rather!than!the!Arctic.!With!a!greater!ability!to!reach!deeperLliving!polychaete!worms!in!the!harshest!of!times,!in!winter!longLbilled!females!have!been!shown!to!stay!put!even!during!cold!snaps.!This!now!turns!out!the!wrong!decision!when!food!abundance!is!low,!a!strategy!now!being!eliminated!by!natural!selection!and!reducing!body!size.!!(

Elasticity(analysis(in(random(matrices(

!Takenori!Takada!

Graduate!School!of!Environmental!Earth!Science,!Hokkaido!University,!KitaLku,!Sapporo!060L0810,!Japan.!!

Projection!matrix!models!have!been!often!employed!as!an!effective!tool!to!quantify!the!population!dynamics!of!both!trees!and!herbs.!Elasticity!analysis!is!the!most!frequentlyLused!analytical!method,!and!Franco!and!Silvertown!published!a!famous!paper,!where!they!mapped!elasticity!vectors!of!survival,!growth!and!fecundity!for!102!plant!species!in!triangle!simplex!and!found!that!they!are!located!in!specific!regions!depending!on!groups!of!plants!with!different!life!history!and!growth!forms.!

To!understand!and!clarify!why!they!are!located!in!specific!regions,!we!constructed!four!types!of!random!matrices.!4!by!4!random!matrices!is!compose!of!two!parts:!fecundity!and!transition!probabilities!from!a!stage!to!another.!The!distribution!of!fecundities!is!a!Poisson!distribution!with!the!average!p.!The!transition!probabilities!range!from!zero!to!one,!whose!row!sums!are!less!than!1.!The!elasticities!for!survival,!growth!and!fecundities!are!calculated!using!3000!random!matrices!and!the!elasticity!vectors!are!plotted!in!the!triangle!map.!Four!types!of!matrices!are!as!follows:!(1)!random!matrices!with!no!zeroLelement,!(2)!random!matrices!with!no!zeroLelement!and!the!survival!probabilities!increase!as!individuals!grow,!(3)!random!matrices!which!have!nonLzero!elements!only!on!diagonal!and!subLdiagonal!positions,!(4)!random!matrices!which!have!nonLzero!elements!only!on!diagonal!and!subLdiagonal!positions!and!the!survival!probabilities!increase!as!individuals!grow.!The!results!are:!(a)!distribution!of!elasticity!vectors!moves!to!upperLleft!region!of!the!triangle!map!as!the!average!of!fecundity!increases.!(b)!In!the!third!and!fourth!type!of!random!matrices,!the!distribution!is!located!on!a!line!whose!slope!is!equal!to!46!degrees.!(3)!There!is!no!elasticity!vectors!in!the!bottom!half!of!the!triangle!map.!eLmail:[email protected]!!(

Equilibria(in(a(population(with(ageOspecific(behaviors(

(

Stefano!Giaimoa,!XiangLYi!Lia,!Arne!Traulsena,!Annette!Baudischb!a!Department!of!Evolutionary!Theory,!Max!Planck!Institute!for!Evolutionary!Biology!b!University!of!Southern!Denmark,!Biology!Department,!MaxLPlanck!Odense!Center!on!the!Biodemography!of!Aging!!

Models!in!evolutionary!game!theory!usually!lack!age!structure.!Yet!these!models!are!often!meant!to!capture!biological!situations!in!which!age!structure!is!present.!The!underlying!assumption!is!that!the!effect!described!by!the!model!is!independent!of!age!structure.!Here,!we!challenge!this!simplification!and!infer!whether!the!equilibrium!states!of!a!gameLtheoretic!model!without!age!structure!are!found!again!in!a!comparable!model!with!age!structure.!We!analyze!both!polymorphic!and!monomorphic!equilibria.!We!show!that!life!histories!can!play!a!crucial!role!in!these!models.!When!the!life!histories!among!competing!types!are!identical,!the!equilibria!of!the!unstructured!model!are!surely!recovered.!There!is!no!such!a!guarantee,!however,!when!life!histories!are!allowed!to!differ.!In!particular,!we!provide!examples!for!social!dilemmas!where!mutually!beneficial!cooperation!is!undermined!by!oneLsided!defection.!Under!life!histories!with!different!rates!of!aging,!cooperation!and!defection!can!coexist,!or!cooperation!cannot!be!invaded,!even!if!encounters!are!random!and!the!population!is!infinite.!! !


! 32!

!!Lifespan(Equality(and(Life(Expectancy(in(Humans(and(Other(Primates((

!Fernando!Colchero,!et!al.,!Annette!Baudisch,!Susan!C.!Alberts,!and!James!W.!Vaupel!!!!

People!are!living!longer!on!average,!but!how!equally!is!this!benefit!distributed!in!a!population?!Social,!economic!and!health!inequality!result!in!lifespan!inequality.!Here!we!reveal!a!lifespan!continuum!along!which!lifespan!equality!increases!in!lockstep!with!average!lifespan,!across!millions!of!years!of!primate!evolution!and!hundreds!of!years!of!human!social!progress.!We!suggest!that!this!remarkable!relationship!drives!and!is!driven!by!enhanced!social!cooperation.!We!hypothesize!that!longer!lives!and!greater!overlap!between!individuals'!lives!promote!increasing!levels!of!knowledge!transfer!and!innovation.!We!also!reveal!that,!across!the!primate!order!and!throughout!human!history,!females!tend!to!have!longer!and!less!variable!lifespans!than!males,!suggesting!deep!evolutionary!roots!to!the!male!disadvantage.!For!humans!this!male!disadvantage!has!grown!exceptionally!large!in!recent!decades,!indicating!social!influences!that!amplify!the!evolutionary!pattern.!Our!findings!open!new!directions!for!research!on!longevity,!ageing!and!sociality.!!

Poster(Presentations(

! 33!

!

Poster!presentations!

For)posters)that)do)not)have)an)abstract)here,)the)abstracts)can)be)found)in)the)contributed)papers)section.)You)can)look)up)the)corresponding)lightning)talk)in)the)program.)!Past(and(Future(Trends(of(Differentials(in(Life(Expectancy(by(Educational(Attainment:(the(Case(of(Egypt!!Huda!Alkitkat!1Ph.D.!in!Biostatistics!and!Demography!!

Life!expectancy!at!birth!is!defined!as!the!average!number!of!years!of!life!expected!by!a!hypothetical!cohort!of!individuals!who!would!be!subject!during!all!their!lives!to!the!mortality!rates!of!a!given!period.!It!is!expressed!as! years.! According! to! the! UN! estimations,! the! average! life! expectancy! at! birth! of! the! global! population!increased!over!the!past!two!decades!from!64.54!years!during!the!period!(1990L1995)!to!70.47!years!during!the!period!(2010L2015)!which!means!that!about!six!more!years!were!added!to!the!average!number!of!years!that!a!newborn!is!expected!to!live.!!This!is!due!to!a!number!of!factors;!one!of!them!is!the!reduction!in!infant!mortality!rates!between!the!same!periods!from!63!deaths!per!1000!live!births!to!36!deaths!per!1000.!

Infant!mortality! rates!differ! according! to! the! socioeconomic! characteristics,! one!of! them! is! the!mother’s!educational!attainment.!Mortality!is!generally!inversely!related!to!mother’s!education!and!mortality!risks!tend!to!be!lower!among!infants!of!educated!mother.!Consequently,! it! is!assumed!that!this!will!be!reflected!in!the!life!expectancy!at!birth.!!

!This!paper!aims!to!study!past!trends!of!infant!mortality!rates!by!sex!and!mother’s!educational!attainment!and!to!estimate!past!and!future!trends!of!the!differentials! in! life!expectancy!at!birth!by!sex!and!educational!attainment!in!Egypt!by!using!data!from!the!EDHS!survey.!

!!Composite(temporal(strategies(in(pathogen(evolution:(balancing(invasion(and(persistence

*(

!Yael!ArtzyLRandrup1!&!Mercedes!Pascual2!

!1!!Theoretical!Ecology!group,!Institute!for!Biodiversity!and!Ecosystem!Dynamics,!University!of!Amsterdam,!eLmail:[email protected]!2!!Department!of!Ecology!and!Evolutionary!Biology!and!Howard!Hughes!Medical!Institute,!University!of!Michigan,!and!Santa!Fe!Institute!

(

There!is!ongoing!interest!in!the!conditions!that!favor!the!evolution!of!acute!highly!transmissible!infections!in!contrast!to!chronic!ones.!Typically,!studies!consider!the!evolution!of!a!pathogens!trait!as!though!it!is!constant!over!the!lifetime!of!an!infection.!However,!for!many!pathogens!different!traits!may!vary!over!this!course.!We!address!here!the!evolution!of!temporal!patterns!in!limited!host!population!sizes,!where!a!tradeLoff!between!invasion!and!persistence!can!arise.!This!is!of!particular!relevance!to!questions!on!the!evolution!of!acuteness!and!chronicity!of!different!diseases.!In!this!study!we!ask!whether!population!dynamics!of!transmission!at!the!betweenLhost!population!level!could!potentially!lead!pathogen!adaptation!to!favor!temporal!strategies!taking!place!during!the!course!of!infection!on!the!within!host!level.!To!do!this,!we!consider!infections!to!be!composed!of!multiple!stages,!and!we!allow!each!of!these!to!evolve!independently!under!a!transmission–duration!tradeLoff.!Importantly,!we!only!consider!here!selection!taking!place!on!the!betweenLhost!level!and!examine!the!balance!of!invasion!and!persistence!(i.e.,!maximizing!replication!vs.!minimizing!vulnerability!to!extinction)!with!several!fitnessLrelated!measures.!We!find!that!a!composite!strategy!that!is!ordered!in!time!can!confer!higher!fitness!than!any!single,!constant,!strategy.!We!discuss!the!relevance!of!these!results!for!the!ordered!expression!of!var!genes!in!Plasmodium%falciparum,!as!well!as!for!infections!that!characteristically!have!several!stages!as!in!some!bacterial!pathogens.!!!!!


! 34!

Contrasting(patterns(of(environmental(fluctuation(contribute(to(divergent(life(histories(among(amphibian(

populations(

(

Hugo!Cayuela¹!²,!Dragan!Arsovski!²,!JeanLMarc!Thirion³,!Eric!Bonnaire4,!Julian!Pichenot5,!Sylvain!Boitaud¹,!AnneLLyse!Brison5,!Claude!Miaud²,!Pierre!Joly¹!Aurélien!Besnard²!!!¹UMR!5023!LEHNA,!Laboratoire!d’Ecologie!des!Hydrosystèmes!Naturels!et!Anthropisés,!69100!Villeurbanne,!France!²UMR!5175!CEFE,!Laboratoire!d’Ecologie!et!de!Biogéographie!des!Vertébrés!(EPHE),!1919!route!de!Mende,!FL34293!Montpellier,!France!³OBIOS!–!Objectifs!BIOdiversitéS,!17250!PontLl'AbbéLd'Arnoult,!France!4Office!National!des!Forêts,!Agence!de!Verdun,!55100!Verdun,!France!5CERFE,!Centre!de!Recherche!et!Formation!en!EcoLéthologie,!08240!BoultLauxLBois,!France!!(

Because!it!modulates!the!fitness!returns!of!possible!options!of!energy!expenditure!at!each!ontogenetic!stage,!environmental!stochasticity!is!usually!considered!a!selective!force!in!driving!or!constraining!possible!life!histories.!Divergent!regimes!of!environmental!fluctuation!experienced!by!populations!are!expected!to!generate!differences!in!the!resource!allocation!schedule!between!survival!and!reproductive!effort!and!outputs.!To!our!knowledge,!no!study!has!previously!examined!how!different!regimes!of!stochastic!variation!in!environmental!conditions!could!result!in!changes!in!both!the!temporal!variation!and!mean!of!demographic!parameters,!which!could!then!lead!to!intraLspecific!variation!along!the!slow–fast!continuum!of!life!history!tactics.!In!this!study,!we!examined!how!contrasting!levels!of!stochastic!environmental!variation!affect!life!history!traits!among!different!populations!of!a!slow!species.!We!predicted!that!increased!environmental!stochasticity!would!increase!yearly!variation!and!decrease!mean!survival!at!ontogenetic!stages.!We!also!predicted!that!increased!environmental!stochasticity!would!make!female!breeding!propensity!more!variable!from!year!to!year.!In!parallel,!we!postulated!that!offspring!productivity!would!also!be!more!variable!from!year!to!year,!but!higher!on!average!when!a!population!experienced!increased!environmental!stochasticity.!Accordingly,!we!predicted!that!increased!environmental!stochasticity!could!speed!up!the!life!history!cycle.!To!investigate!these!hypotheses,!we!used!capture–recapture!data!collected!on!five!populations!of!a!longLlived!amphibian!(Bombina%variegata)!experiencing!two!distinct!levels!of!stochastic!environmental!variation:!(i)!constant!availability!of!breeding!sites!in!space!and!time!(‘predictable!environment’),!and!(ii)!variable!spatioLtemporal!availability!of!breeding!sites!(‘unpredictable!environment’).!We!found!that!female!breeding!propensity!varied!more!from!year!to!year!in!unpredictable!than!in!predictable!environments.!Although!females!in!unpredictable!environments!produced!on!average!more!viable!offspring!per!year,!offspring!production!was!more!variable!between!years.!Survival!at!each!ontogenetic!stage!was!slightly!lower!and!varied!significantly!more!from!year!to!year!in!unpredictable!environments.!Taken!together,!these!results!confirm!that!increased!environmental!stochasticity!can!modify!the!resource!allocation!schedule!between!survival!and!reproductive!effort!and!outputs!and!may!lead!to!intraLspecific!variation!along!the!slow–fast!continuum!of!life!history!tactics.%!Patterns(in(frailty(in(human(populations;(a(comparison(between(countries,(sexes(and(periods(

(

Nienke!Hartemink,!Trifon!Missov!and!Hal!Caswell!IBED,!University!of!Amsterdam,!The!Netherlands!Max!Planck!Institute!for!Demographic!Research,!Rostock,!Germany!!

Individual!heterogeneity!or!frailty!in!mortality!risk!is!an!important!concept!in!human!demography!and!it!is!now!also!being!used!in!models!for!animal!longevity.!In!this!study,!we!illustrate!a!method!to!estimate!frailty,!and!its!contribution!to!variance!in!longevity,!in!human!populations,!and!compare!frailty!estimates!between!countries,!between!the!two!sexes!and!between!different!periods.!

We!estimate!the!GammaLGompertz!(GG)!and!GammaLGompertzLMakeham!(GGM)!parameters!for!each!birth!cohort,!based!on!cohort!mortality!data.!Incorporating!these!parameter!values!in!a!vecLpermutation!matrix!model!(Caswell!2014,!Demographic!Research)!allows!us!to!calculate!longevity!statistics.!!The!variance!in!longevity!can!be!decomposed!into!two!parts:!one!part!attributable!to!heterogeneity!and!one!to!individual!stochasticity.!!We!do!this!for!three!European!countries!with!good!mortality!records:!France,!Italy!and!Sweden.!The!analysis!was!done!per!birth!cohort,!for!several!starting!ages!(as!GG!is!known!to!perform!well!at!higher!ages!but!not!at!capturing!mortality!patterns!at!young!ages)!and!for!the!two!sexes!separately.!!


! 35!

!The!proportion!of!variance!due!to!heterogeneity!is!higher!when!incorporating!GGM!model!derived!parameters!in!the!matrix!model!than!when!using!GG!model!derived!estimates.!When!looking!at!the!temporal!pattern,!it!is!particularly!high!for!cohorts!that!experience!a!mortality!crisis!at!the!start!of!the!analysis,!such!as!males!in!France!and!Italy!when!starting!the!analysis!at!1914L1919,!the!period!of!WWI!and!the!Spanish!flu.!This!suggests!that!the!effect!of!mortality!crisis!needs!to!be!taken!into!account!in!this!type!of!analysis.!!

Poor!identifiability!of!the!GGM!and!GG!models!makes!it!hard!to!draw!conclusions!on!absolute!levels!of!heterogeneity,!which!is!why!we!focus!on!comparing!the!same!type!of!models!and!on!identifying!patterns!and!trends!in!time!and!between!countries.!!!!Climate(Change(Alters(Mating(Strategies(and(Social(Systems(Among(Rural(African(Pastoralists!Ashley!Hazel1,2!&!James!Holland!Jones2,3!1Department!of!Anthropology,!Stanford!University;!2Woods!Institute!for!the!Environment,!Stanford!University;!3Department!of!Environmental!Earth!Systems!Science!!

The!Himba,!a!semiLnomadic,!agroLpastoralist!population!in!northwestern!Namibia,!like!most!African!pastoralists,!live!in!arid!landscapes!that!are!vulnerable!to!drought.!To!sustain!livestock!herds!under!these!conditions,!pastoralists!employ!a!variety!of!strategies!to!adapt!to!periods!of!extreme!weather,!including!transhumance!movement,!dispersal,!supplemental!subsistence!(e.g.!smallLscale!cultivation),!or!seasonal!labor!migration.!Mobility!determines!sexual!partner!access!and!influences!cultural!norms!about!relationship!dynamics,!fertility,!and!family!structure.!Concurrent!sexual!partnerships!are!central!to!the!highLmobility!adaptations!of!the!Himba.!Concurrency!gives!women!freedom!to!choose!sexual!partners!for!emotional!support!or!resource!access,!even!after!entering!into!arranged!marriage.!However,!concurrency!also!results!in!high!paternity!uncertainty.!To!bypass!proof!of!paternity!and!relieve!familial!tensions,!the!Himba!use!matrilineal!descent!for!livestock!inheritance.!

As!climate!change!increases!the!frequency!and!length!of!droughts!in!arid!environments,!strategies!that!used!to!be!temporary!adaptations!for!optimizing!resource!access!during!annual!dry!seasons!or!unforeseen!droughts!could!become!permanent!solutions!for!life!in!an!altered!climate,!in!turn!affecting!fertility,!parental!investment,!and!lineage!persistence.!We!have!observed!early!signs!of!behavioral!change!in!response!to!a!15Lyear!drought!among!the!Himba.!

We!employ!stochastic!dynamic!programming!to!evaluate!optimal!mobility!strategies!for!different!objective!functions!relating!to!economic!returns,!fertility,!and!longLterm!lineage!persistence!under!regimes!of!ecological!volatility.!Models!are!parameterized!using!data!collected!from!ongoing!work!among!Himba.!

Models!predict!that!as!nonLherding!strategies!become!optimal!for!an!increasing!proportion!of!the!population,!overall!mobility!will!decrease.!Furthermore,!for!most!women,!concurrency!will!decrease,!but!will!increase!for!a!minority!of!unmarried!women.!We!expect!that!the!resulting!increased!paternity!certainty!may!shift!inheritance!from!the!matriline!to!the!patriline.!!A(metaOanalysis(into(the(declines(of(fecundity(and(parental(investment(with(increasing(maternal(age(

!Edward!IvimeyLCook!and!Jacob!Moorad!University!of!Edinburgh!!

Numerous!studies!have!shown!that!fecundity!and!parental!investment!decline!with!increasing!maternal!age.!However,!little!is!known!about!how!senescence!in!maternal!investment!and!fecundity!differ.!A!recent!theoretical!model!predicts!a!faster!decline!of!maternal!investment!with!age!compared!to!fecundity,!but!results!from!empirical!studies!vary!across!the!animal!kingdom.!Therefore,!a!metaLanalysis!was!conducted!to!compare!the!senescence!of!fecundity!and!investment!when!maternal!age!increased.!A!total!of!9000!papers!were!initially!found!after!an!extensive!literature!search!involving!specific!strings!within!Scopus!and!Web!of!Knowledge.!These!papers!were!then!assessed!and!reduced!to!include!only!those!that!contained!a!measure!of!fecundity!and!investment!over!a!mother’s!lifespan.!The!patterns!of!senescence!in!maternal!fecundity!and!investment!were!then!compared!across!taxa!and!with!the!theoretical!predictions!implied!by!the!model.!! !


! 36!

!CoOvariation(patterns(between(survival(and(reproductive(effort(at(an(intraOannual(scale(in(a(shortOlived(

species,(the(Siberian(chipmunk((Tamias)sibiricus)(

Christie!Le!Coeur,!JeanLLouis!Chapuis!and!Alexandre!Robert!

Centre!d’Ecologie!et!des!Sciences!de!la!Conservation!(CESCO!UMR!7204),!Sorbonne!Universités,!!MNHN,!CNRS,!UPMC,!CP51,!57!rue!Cuvier,!75005,!Paris,!France!

Positive!or!negative!coLvariations!among!fitness!components!are!central!to!lifeLhistory!theory.!From!the!resource!allocation!principle,!current!reproduction!is!predicted!to!be!costly!in!terms!of!future!fecundity!or!survival,!engendering!negative!coLvariation!among!fitness!components.!In!contrast,!demographic!heterogeneity!can!lead!to!positive!coLvariations!among!lifeLhistory!traits.!In!shortLlived!species!(fast!lifeLhistory!strategy),!individuals!are!expected!to!allocate!a!high!reproductive!effort!at!each!of!their!few!reproductive!occasions,!involving!subsequent,!shortLterm!costs.!Highlighting!such!variation!and!covariation!patterns!in!natural!populations!is!challenging.!

In!this!context,!we!studied!coLvariation!patterns!between!survival!and!reproductive!effort!at!an!intraLannual!time!scale!in!a!shortLlived!species,!characterized!by!two!reproductive!seasons!(in!early!spring!and!early!summer),!associated!with!different!resource!availabilities!and!a!hibernation!period.!The!Siberian!chipmunk!(Tamias%sibiricus)!is!a!small!mammal!introduced!to!an!oakLhornbeam!forest!in!France,!the!forest!of!Sénart!(Essonne,!France).!This!introduced!population!has!been!monitored!since!2004!by!captureLmark–recapture!procedure.!The!relationship!between!reproduction!effort!and!survival!was!investigated!at!a!seasonal!scale!(summer!and!winter!survival!intervals)!based!on!the!lactation!status!of!adult!females!(lactating/nonLlactating,!Nfemales=262)!at!both!breeding!seasons,!using!multiLstate!and!multiLevent!models!(ELSURGE!software).!The!results!revealed!!(i)!no!evidence!of!negative!effect!of!the!reproductive!effort!in!the!season!immediately!preceding!the!onset!of!hibernation!(ie,!summer!reproduction)!on!subsequent!overwinter!survival,!but!(ii)!strong!evidence!of!negative!delayed!effect!of!early!reproductive!effort!(spring!reproduction)!on!the!following!winter!survival.!Our!results!suggest!the!existence!of!a!cost!of!reproduction!modulated!by!the!variation!of!resource!availability!at!an!intraLannual!scale.!!A(global(estimate(of(spread(velocity(as(a(function(of(plant(species(life(history!!Nicky!Lustenhouwer*,!Emily!V.!Moran†!and!Jonathan!M.!Levine*!

!*!Institute!of!Integrative!Biology,!ETH!Zurich,!Universitaetstrasse!16,!8092!Zurich,!Switzerland!†!School!of!Natural!Sciences,!UC!Merced,!5200!Lake!Rd,!Merced,!CA!95343,!USA!!

Population!spread!lies!at!the!heart!of!two!pressing!concerns!to!ecologists:!the!growing!threat!of!biological!invasions!and!range!shifts!of!native!species!in!response!to!climate!change.!Theory!predicts!that!species!with!high!dispersal!and!fecundity!and!a!short!time!to!maturity!should!have!the!highest!capacity!to!spread.!However,!empirical!studies!of!spreading!plant!species!have!reported!mixed!results!as!to!which!plant!life!form!has!spread!fastest!in!the!past.!This!discrepancy!could!reflect!a!biological!reality:!life!history!traits!that!promote!spread!may!trade!off!with!each!other,!which!makes!it!unclear!which!plant!life!form!L!if!any!L!should!be!expected!to!spread!the!fastest.!Here!we!show!that!life!history!tradeLoffs!equalize!spread!velocities!across!plant!taxa,!ranging!from!annual!herbs!to!trees.!In!a!comparative!analysis!of!demographic!and!dispersal!measures!of!78!plant!species,!we!found!tradeLoffs!across!species!between!early!maturation!and!both!far!dispersal!and!high!fecundity.!We!then!input!these!values!into!a!stochastic!model!of!spread!to!predict!which!species!would!spread!the!fastest.!Similar!rates!of!spread!were!estimated!across!a!gradient!from!early!maturing!species!with!poor!dispersal!and!low!fecundity,!such!as!annual!plants,!to!late!maturing!species!with!high!dispersal!and!fecundity,!such!as!trees.!However,!variance!in!spread!velocity!was!high,!with!the!highest!spread!velocities!predicted!for!early!maturing!species!with!strong!dispersal!capacities.!Detailed!dispersal!data!are!therefore!required!for!species!level!predictions!of!the!spread!velocity.!Considering!multiple!life!history!traits!simultaneously,!our!results!demonstrate!that!remarkably!different!plant!life!forms!may!reach!similar!spread!velocities.!! !


! 37!

!!Forest(dynamics(in(the(context(of(the(slowOfast(continuum(

Sean!M.!McMahon!!Smithsonian!Institution!!

LiveLfast!die!young.!!Slow!and!steady!winds!the!race.!!These!cliche’s!define!the!extremes!of!the!slowLfast!continuum.!!The!negative!relationship!between!growth!and!mortality!for!forest!tree!species!has!been!invoked!as!a!method!of!fostering!coexistence,!and!yet!careful!analysis!of!demographic!vital!rates!and!their!integration!into!populationLscale!metrics!(such!as!passage!time!or!longevity)!has!not!been!pursued!in!forest!evolutionary!ecology.!!Here!I!use!a!caseLstudy!of!a!central!african!forest!to!demonstrate!both!the!observed!patterns!of!vital!rates!and!how!their!integration!in!an!IPM!framework!can!illuminate!biogeographical!patterns!of!forest!change!over!time.!!Further,!there!is!evidence!that!‘darwinian!monsters’,!species!that!exist!off!of!the!tradeLoff!axis!of!the!slowLfast!continuum,!play!a!unique!but!important!role!in!many!forests.!!This!has!implications!for!evolutionary!theory!of!coexistence,!as!well!as!ecological!models!of!biome!change!under!climate!change.!!(

MorphOspecific(changes(in(buzzard(vital(rates(increase(the(fitness(of(extreme(morphs(and(explain(rapid(

population(growth(

Astrid!Potiek,!Behavioral!Ecology,!Bielefeld!University,!!(

Phenotypic!variation!often!results!in!fitness!differences.!These!fitness!differences!may!change!over!time!as!a!result!of!changing!selection!pressures.!Understanding!the!temporal!dynamics!of!fitness!differences!is!vital!for!population!dynamics!and!especially!longLterm!growth!trajectories.!We!measured!annual!and!lifetime!fertility!and!calculated!survival!using!markLrecapture!data!in!a!German!population!of!common!buzzard!Buteo%buteo,!over!the!last!25!years.!This!species!shows!a!genetically!determined!plumage!polymorphism,!comprising!three!colour!morphs.!Survival!increased!most!strongly!for!the!extreme!morphs,!whereas!fertility!stayed!relatively!constant!in!all!morphs.!Using!morphL!and!ageLspecific!matrix!models,!we!show!that!the!relative!contribution!of!three!morphs!to!population!growth!changed!over!time.!We!calculated!the!net!reproductive!output!R0,!which!is!a!measure!for!fitness!giving!the!expected!future!offspring!for!a!certain!stage!and!phenotype.!Whereas!previously!only!the!intermediate!morph!had!a!R0Lvalue!above!one,!more!recent!data!indicate!that!both!extreme!morphs!now!contribute!to!population!growth!as!well.!Moreover,!we!show!that!changed!demographic!parameters!explain!the!rapid!population!growth!relatively!well,!and!that!net!immigration!seems!to!play!a!minor!role.!Elasticity!values!suggest!that!increased!survival!seems!to!be!the!main!driver!of!population!growth,!with!stronger!increases!in!the!extreme!morphs.!Climate!change!seems!to!be!the!cause!for!this!increase!in!survival,!and!hence!for!the!change!in!relative!fitness.!Extreme!morphs!seem!to!experience!weaker!selection!pressure!during!winter!than!previously,!resulting!in!smaller!fitness!differences.!This!shows!how!a!changed!selection!pressure!as!a!result!of!mild!winters!can!change!the!relative!fitness!of!different!phenotypes.!!!Age,(sex(and(the(cost(of(reproduction(in(an(extremely(longOlived(herb(

!Danielle!Sherman!(Department!of!Biology,!MaxLPlanck!Odense!Centre,!MaxO),!Johan!P.!Dahlgren!(Department!of!Biology,!MaxLPlanck!Odense!Centre,!MaxO),!Johan!Ehrlén!(Department!of!Botany,!Stockholm!University,!SEL10691!Stockholm,!Sweden)!and!Maria!B.!Garcia!(Instituto!Pirenaico!de!Ecologı´a!(CSIC),!Apdo.!13034,!50080!Zaragoza,!Spain)!!We!report!on!our!study!of!life!history!tradeLoffs!in!the!longLlived!dioecious!herb!Borderea%pyrenaica!with!the!specific!aim!of!understanding!why!this!species!seemingly!does!not!show!demographic!senescence.!We!also!examine!possible!sex!differences!in!tradeLoff!within!this!species.!Observational!and!experimental!data!were!collected!by!Maria!Garcia!over!a!fiveLyear!period!in!the!Spanish!Pyrenees!and!analysed!using!generalised!linear!models.!We!discuss!how!observed!patterns!of!the!costs!of!reproduction!may!contribute!to!the!lack!of!decrease!in!reproductive!value!with!increasing!age!observed!in!both!females!and!males.!!! !


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!The(impact(of(immigration(on(longOterm(and(transient(population(dynamics(

Anna!Zincenko!Southhampton!University,!UK!!We!propose!the!simple!matrix!female!population!model!that!includes!emigration!and!immigration.!In!this!model!we!aggregate!emigration!with!mortality!and!consider!immigration!as!a!given!external!inflow.!The!main!assumption!is!that!the!children!of!migrants!born!in!Germany!belong!to!the!native!population.!This!means!that!their!demographic!parameters!coincide!with!those!of!native!population.!The!data!was!obtained!from!Destatis,!data!reconstruction!was!performed!to!disaggregate!the!data!to!the!required!level!of!detail,!especially!with!respect!to!age!groups.!We!have!shown!that!constant!growth!of!migrant!inflow!generates!large!transient!fluctuations!in!the!German!born!and!migrant!born!population.!In!particular,!for!population!of!Germany!the!critical!rate!of!growth!lambda!of!immigration!flow!is!calculated!at!which!the!proportion!of!migrants!in!Germany!reaches!50%!after!some!time!period.!This!critical!rate!does!not!exceed!1.03!in!other!words!does!not!exceed!the!growth!of!3%!per!year,!and!its!value!is!not!sensitive!to!some!reasonable!changes!to!the!age!structure!of!migrant!inflow.!On!the!other!hand,!the!time!for!which!the!fraction!of!migrants!reaches!50%,!for!greater!than!the!critical!value,!is!very!sensitive!to!the!value!of!lambda.!In!methodological!terms,!the!proposed!model!allowed!calculating!transient!fluctuation!of!age!structure!of!the!population,!and!estimating!the!time!of!stabilisation!in!the!presence!of!migration.!Further!work!includes!relaxing!the!assumptions!on!the!constancy!of!vital!rates!and!migration!streams,!and!carrying!out!a!more!inLdepth!sensitivity!analysis.!

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