The evolution of insect life history strategies in a social context [Elektronische Ressource] / vorgelegt von Oliver Mitesser
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The evolution of insect life history strategies in a social context [Elektronische Ressource] / vorgelegt von Oliver Mitesser

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THE EVOLUTION OF INSECTLIFE HISTORY STRATEGIESIN A SOCIAL CONTEXTDISSERTATION ZUR ERLANGUNG DES NATURWISSENSCHAFTLICHENDOKTORGRADES DER BAYERISCHEN JULIUS-MAXIMILIANS-UNIVERSITÄTWÜRZBURGVORGELEGT VONOLIVER MITESSERAUSSCHWEINFURTWÜRZBURG 2007Eingereichtam:22.Dezember2006MitgliederderPrüfungskommission:Vorsitzender:Prof.Dr.RolandBenzErstgutachter:Prof.Dr.HansJoachimPoethkeZweitgutachter:Prof.Dr.ErhardStrohmTagdesPromotionskolloquiums:02.Mai2007Doktorurkundeausgehändigtam:..........................Allcold bloodedanimalsspendanunexpectedlylargeproportionoftheirtimedoingnothingatall,oratanyrate,nothinginparticular.C.Eltonin"Animalecology",1927FürUschiTableofContentsTableofContents 71 Introduction 91.1 Colonycycleinannual,primitivelyeusocialbeesandwasps . . . . . 101.2 Optimisationmodelsinevolutionaryecology . . . . . . . . . . . . . 141.3 Dynamicanalysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211.4 Evolutionaryanalysisofthecolonycycle . . . . . . . . . . . . . . . 271.5 Scopeandoutlineofthethesis . . . . . . . . . . . . . . . . . . . . . 32PartI:Activityversusinactivity 352 An ESS model for the evolution of dimorphic development strategies inthebutterflyMaculinearebeli,asocialparasiteofMyrmicaantcolonies 372.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 392.2 Modelandresults . . . . . . . . . . . . . . . . . . . . . . . . . . . . 422.3 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . .

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Publié le 01 janvier 2007
Nombre de lectures 17
Langue English
Poids de l'ouvrage 3 Mo

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THE EVOLUTION OF INSECT
LIFE HISTORY STRATEGIES
IN A SOCIAL CONTEXT
DISSERTATION ZUR ERLANGUNG DES NATURWISSENSCHAFTLICHEN
DOKTORGRADES DER BAYERISCHEN JULIUS-MAXIMILIANS-UNIVERSITÄT
WÜRZBURG
VORGELEGT VON
OLIVER MITESSER
AUS
SCHWEINFURT
WÜRZBURG 2007Eingereichtam:22.Dezember2006
MitgliederderPrüfungskommission:
Vorsitzender:Prof.Dr.RolandBenz
Erstgutachter:Prof.Dr.HansJoachimPoethke
Zweitgutachter:Prof.Dr.ErhardStrohm
TagdesPromotionskolloquiums:02.Mai2007
Doktorurkundeausgehändigtam:..........................Allcold bloodedanimalsspendanunexpectedlylarge
proportionoftheirtimedoingnothingatall,oratanyrate,nothinginparticular.
C.Eltonin"Animalecology",1927FürUschiTableofContents
TableofContents 7
1 Introduction 9
1.1 Colonycycleinannual,primitivelyeusocialbeesandwasps . . . . . 10
1.2 Optimisationmodelsinevolutionaryecology . . . . . . . . . . . . . 14
1.3 Dynamicanalysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
1.4 Evolutionaryanalysisofthecolonycycle . . . . . . . . . . . . . . . 27
1.5 Scopeandoutlineofthethesis . . . . . . . . . . . . . . . . . . . . . 32
PartI:Activityversusinactivity 35
2 An ESS model for the evolution of dimorphic development strategies in
thebutterflyMaculinearebeli,asocialparasiteofMyrmicaantcolonies 37
2.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
2.2 Modelandresults . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
2.3 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
2.4 Appendix:Mathematicalderivations . . . . . . . . . . . . . . . . . . 55
3 The influence of soil temperature on the nesting cycle of the halictid bee
Lasioglossummalachurum 67
3.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
3.2 Materialandmethods . . . . . . . . . . . . . . . . . . . . . . . . . . 70
3.3 Results. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
3.4 Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
4 The evolution of activity breaks in the nest cycle of annual eusocial bees:
amodelofdelayedexponentialgrowth 83
4.1 Background . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85
4.2 Results. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87
4.3 Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94PartII:Workersversussexuals 99
5 Depletion of fat reserves during hibernation and nest establishment in
foundressqueensofthehalictidLasioglossummalachurum 101
5.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103
5.2 Materialsandmethods . . . . . . . . . . . . . . . . . . . . . . . . . 105
5.3 Results. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107
5.4 Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109
6 Optimal investment allocation in primitively eusocial bees: a balance
modelbasedonresourcelimitationofthequeen 113
6.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 115
6.2 Asimpleinvestmentbalancemodel . . . . . . . . . . . . . . . . . . 117
6.3 Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122
7 Adaptivedynamicresourceallocationinannualeusocialinsects:Environ
mentalvariationwillnotnecessarilypromotegradedcontrol 127
7.1 Background . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 129
7.2 Results. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 131
7.3 Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139
7.4 Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143
Summary 145
Zusammenfassung 151
References 157
Publications 173
Conferences&Talks 175
CurriculumVitae 177
Danksagung 179
Erklärung 181Chapter1
Introduction
The analysis of life histories is essentially aiming for an understanding of the evolu
tionofthetemporalsequenceofdecisionsbetweenbehaviouralalternativesduringan
organism’s entire life time. Conspicuous behavioural characteristics of an organism’s
life history are transitions between activity and inactivity, such as between diapause
and development, between sleeping and being awake, and plainest life and
death. Certainly choices between other alternatives are of interest, too: the transition
from growth to reproduction, the number and timing of reproductive events, the
number of offspring, or the choice of a suitable habitat for foraging or oviposition.
These and many other characteristics of life vary both between species and between
individuals of the same species and their evolution has been addressed in numerable
lifehistorystudies(Roff,1992;Stearns,1992).
Sociobiology and theory of social evolution expanded classical life history analysis
from solitary to social organisms. Hamilton’s rule (1964) and Triver’s (1971) concept
of reciprocal altruism paved the way for explaining the evolution of social behaviours
such as altruism, aggression, brood care and nourishment. However, since then the
relevance of kin selection in the context of social evolution has gradually been down
played and partly been substituted by explanations based on ecological factors. The
temporal variability in life history decisions of social insects due to ecological and
demographic factors has been investigated only in a few classical modelling studies
(Macevicz & Oster, 1976; Oster & Wilson, 1978). As in evolutionary life history
analysis in general, the mathematical formalisation of the biological system and the
behavioural alternatives played a key role in these studies. Astonishingly, the original
approach has hardly been continued since then (but see Karsai et al., 1996; Beekman
etal.,1998a).
This thesis extends the classical work of Macevicz and Oster (1976, expanded by
OsterandWilson,1978).Itfocusesontheevolutionofdynamicbehaviouralpatternsin
socialinsectsasaconsequenceofoptimalallocationofenergyandtimeresources.Itis10 Chapter1–Introduction
basedondetailedempiricalobservationsinthemodelspeciesLasioglossummalachu
rum(Halictidae;Hymenoptera,seefigure1.1).Themaintopicsofthisthesisareopti
misationmodelsforeusociallifehistories,temporalvariationinlifehistorydecisions,
and annual colony cycles of eusocial insects. These aspects will be introduced in the
nextsections.
1.1 COLONY CYCLE IN ANNUAL, PRIMITIVELY
EUSOCIAL BEES AND WASPS
The best studied groups of insects with primitively eusocial colony organisation and
annual nest cycles are halictid bees (Halictidae; Hymenoptera), vespid wasps (Vespi
dae; Hymenoptera) and bumble bees (Bombini; Apidae; Hymenoptera). Although the
empirical work of our studies focused on the halictid bee Lasioglossum malachurum,
mostofthetheoreticalresultscanalsobeappliedtoanyotherspeciessharingspecific
characteristicsoftheirnestcycleswithhalictids(seealsofigure1.2).Thus,classifying
the species under consideration by their taxonomic subsumption is only possible at a
verycoarselevel.
The halictid bees show nearly all levels of social organisation, from solitary, to
subsocial,communal,quasisocial,semisocial,andprimitivelyeusocialspecies(Mich
ener, 1974; Sakagami, 1974; Michener, 2000). Phylogenetic analyses revealed that
eusocialityinhalictidshasevolvedindependentlyseveraltimesandhasbeenlosteven
more often (Wcislo & Danforth, 1997; Danforth, 1999; Danforth et al., 1999; Dan
forth, 2002; Danforth et al., 2003). Most species have an annual life cycle (due to
annualvariationbetweencoldandwarmordryandwetseason)butthisdoesnotneed
to be so. There is at least one perennial halictid species (Lasioglossum marginatum,
Packer (1991)), and several tropical species do not exhibit a diapause, i.e. nest found
ingoccursthroughoutthewholeyear(Wcislo,1997a).
Vespidwaspsmainlyincludesolitaryandeusocialspecies.Both,paperwasps(Polisti
nae; Vespidae) and yellowjackets (Vespinae; Vespidae) are subfamilies with exclu
sivelyeusocialspecies.Althoughthetypicalcolonycycleisannual,occasionallynests
inshelteredsitesorduringmildwintersarenotabandoned.Thenaperennialcolonycy
clemaybemaintainedbythequeenandtheworkersfromthepreviousyear.Perennial
coloniesaregenerallylargerthanannualcolonies(Spradbery,1973;Ross&Mathews,
1991;Yoshiaki,1993).
Besidethehighlyeusocialhoneybees(Apini)andstinglessbees(Meliponini)theApi
daefamilyalsoincludesseveraltribiandgenerawithspeciesexclusively(bumblebees
(Bombini, see Goulson, 2003), orchid bees (Euglossini, see Dressler, 1982)) or partly
primitively eusocial (carpenter bees (Xylocopa, e.g. Steen & Schwarz, 2000)), allo

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