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The Role of Env 1 Ronmental Heterogeneity in the Evolution Of f THE ROLE OF ENV 1 RONMENTAL HETEROGENEITY IN THE EVOLUTION OF J..IFE HISTORY STRATEGIES OF THE STRIPED GROUND CRICKET Michael James Bradford Department of Biology McGill University, Montréal December 1991 A thesis submitted to the Faculty of Graduate Studies and Research in partial fulfilment of the requirements for the degree of Doctor of Philosophy. @ Michael J. Bradford il ABSTRACT r cxamincd the E·ffect of heterogeneity in the thermal environment on the 11[e history o[ the cricket Allonemobius fasciatus. Variation in the lHe cycle was the result of a mixture of phenotypic plasticity and gcnet_ic differentlation in phenology-related traits along a latitudinal cline in glowing season. Females from a partially bivoltine population hllve a conditional life history because the y can adjust the proportion of diapause eggs in accordance with the likelihood that a second generation will grot" and reproduce before winter. The thermal environment is not variable enough to resul t in the evolution of a mllrked bet hedging response, Ils is predicted by the ory . A quantitative genetic analysis of the diapause reaction norm revealed significant heritabilities as weIl as correlations with other traits that could be related to common physiological mechanisms. c iii RESUME J'ai examiné l'effect de l'hétérogénoité do l'Pllvlrollllcmcnl thermique sur la cyclo-biologie du grillon Allonemobius faseiatuR. Une variation du cycle vital issue d'un mélange de plasticHé phénol ypique et la différentiation génétique de traits phénologiqucs [ut rel i 0(' A Ull(' tendance nord-sud influençant la longeur de lél sa i son dt' crot ssnllCC. LN; femelles provenent d'une population partiellement hi volt i llC ont LIllC' cyclo-biologie candi tionelle puisque' elles peuvant ajusLer 1a propol't i 011 des oeufs en diapause suivant la probabilité qu'une scconc\(' !j0néri-ll ion se reprodiuse avec succès avant l'hiver. L'environnement thennic}llC' n'cHI pas assE"Z variable pour favoriser l'évolution d'une sLrnl-égic précisc dC' minimimisation des risques de mortalité, tel que prédit par ln I-hi·orip. Une analyse génétique quantitative a démontré 1 ' exi sl-anc:e d'héritabilit~s significative et de corre]atLons entre dirf~rents trnils reliés à des méchanismes phsio10giques communs. (' iv PREFACE The [aculty of Graduate Studies and Research requires the following statement to be roprinted: "'rhe candidate has the option, subject to the approval of their Department, of including as part of the thesi s the text, or dupl icated published text, of an original paper or papers. Manuscript-style theses must still conform to aIl other requirements explained in the Guidelines Goncerning Thesis Preparation. Additional material (procedural and dosign data as weIl as descriptions of equipment) must be provided in sufficient detail (e.g. 1 in appendices) to allow clear and precise judgcment to be made of the importance and original i ty of the research reported. The thesis should be more than a mere collection of manuscripts published or to be published. 1t must include a generaI abstract, a full introduction and literature review and a final overall conclusion. Connecting texts which provide logical bridges between difEerent manuscripts are usually desirable in the interest of cohesion. "It is acceptable for theses to include, as chapters, authentic copies of papers already published, provided these are duplicated clearly and bound as an integral part of the thesis. ln such instances, connecting texts are mandatory a supplementary explanatory material is always necessary. Photographs or other materials which do not duplicate wall must be included in their original form. "While the inclusion of manuscripts co-authored by the candidate is acceptable. the candidate is required to make an explicit statement ln the thesis of who contributed to such work and to what extent, at supervisors must attest to the accuracy of the c1aims at the Ph. D. Oral Defence. Since the task of the Examiners is made more difficult in these cases, it is in the candidate' s interest to make the responsibili ties of v authors perfectly clear," In accordance with the above. my thesis consi!lts of [0111' manuseripts intended for suhmissi on to scient i fi c joli\' na 1 s, i\ gpnC'I'nl summary has been added to provide an overview of Lhe r tC'l ci. 1I1ld t 0 indieate how the manuscripts an:! linked in wha\ l ùelicvo to hl' H logieal sequence of investigation, AlI chaptcrs deal wi th cl Lf! corollt aspects of a single theme. and this resul ts in some dupl tO.lt i on of t Il(' introduetory material. Nonetheless the introductions, in their C'ntlrcly. provide a thorough literature review. There art' slightly di/fol'C'nl styles used in the presentation of thC' text, in anticipatioll or tl\(' manuscripts being sent to different journals. At the lime of wr i t i 118 (September 1991), only Chapter 2 has been submitted, Lo the journrtl Eeology, AU four chapters are written in the plural voice. llIleI ni 1 will he' co-authored with my thesis supervisor. Dr. Derek Roff, I\llhough Dr, Hon has provided much advice in the design, execut ion and ana1ys i s of 1 hi!> research, his contribution falls wi thin the reillm of Il thes i Il supervisor; the thesis represents largcly my own work, Contributions ta Original Knowledge 1) Although phenotypie plasticity can relieve the nced [or gerll'tic diversification in the adaptation to diverse habi Lat s, l WflS Ilbl e Lo demonstrate in the case of diapause ln Allonemobius EasciaLus, thllt bath plasticity and genetie differentiation are requircd, 2) A detailed investigation of the 1 He history of individuaJ [cmlJlNl 1ead to different conclusions about the role of envi ronmental heterogeneity than would have been made from overa] l popuLttLon vi ( averages. Recause sc] cc t j on ae ts on indi vidual s, this resul t underscorcs the nood to study lifc history variation at the level of the individual. 3) This i s the first use of long-term meteorologicai records ta study li le history evolution for a specifie organism. Al though insect seasonal i ty has becn used as an example in a vadety of theoretical mode} s of ovo] ution in a variable cnvironment. these moclels have rarely been Lested. 'f) The final chapter is one of thE' first investigations of the quanti tat! vc geneties of li c1early adaptive case of phenotypic plasticity. 1 was able ta relate aspects of this plasticity ta different types of selection imposed by the thermal regime, as indicated by my modell1ng work. The patterns of trait correlations were aiso suggestive of a common hormonal mechanism regulating the expression of these plastic traits. 5) Fillally, from a technical perspective, this thesis ls one of the few cxamples of the successful use, in laboratory experiments, of naturally changing photoperiod and tempe rature cycles ta induce the patterns phenotypic variation seen in the field. vi i • ACKNOWLEJ)GEMENTS First and foremost r must thank my thcsis supC'rvlsor. D"rpl, Rofr. for his role in this research. He fj t"st suggcstcd t 11:11 i nSl'cl H 1111 ght hl' a very profitable way of studying t11C' evolutlon of Hfl' hisloriC's. and his guidance throughout has made my ontry into titis fll'ld v(,l"y rewarding. l must also thank Tim Mousseau for inlroduc:illg 1IIl' 10 crickets, for tolerating many hours of quest 1011 1ng, for hl' i nE, encollraging throughollt and providing the tempera turc dat <1 of Cillipler 3. Members of my supervisory commi ttee. Don Kramcr and Gntll(llll He 11 . provided usefui advLce, especiaIIy in the early going. Mnrk JOhllstoll mld Robert McLaughlin commented on drafls of some of t lw chnpt ('l'S. Gi Il)('1'1 Cabana had something to say about near]y evcrythillg lIncl lrmlslaLpc! IIIl' abstract. Rearing crickets takes III11ny hours of work, fllld 1 t hflllk l'mil Guerette, Nathalie Roy. Jackie Farrell and Sharon David fOI" , hpl r III') p. Special thanks to AnnR Chandler, who ran the expel'Î lOc>nl of Clwpll'r ï 011 her spare time. Finally, thanks to aIl the 6th floor people who mmll' my four year stay at McGill fly by. Financial support was provl d('d by 1111 NSERG and Max Bell post-graduate scholarships, as wc 11 as NSlmC operating grants ta D. Roff. This thesis ls dedicated to LOllisC' Fock1!'1- and Anna Bradford. for encouragement. SUprot"t and love 1Il th<.' PlIS! , present and future . .". viii TABLE OF CONTENTS • ABSTRACT. • . .. ii RESUME .............................................. " . i ii PRgFACE .............................. " . ..... • ... ... .. .. iv ACKNOWLEOGEMENTS. .. vii 1.1 ST OF TABLES. .. x LIST OF FIGURES ...................................•............. xi GENgRAL OVERVI EW. 1 GI~PTER 1: Life history variation along a cline in voltinism in the cricket A11onemobius fasciatus. Abstract. 10 Introduction ...................................... Il Methods and Materlals... 13 Results ............................................ 17 Discussion. 31 Conclusions. .. ................................. 36 References. 37 CIIAPTER 2: Bet-hedging and phenotypic plasticity in the diapause strategies of the cricket Al1onemobius fasciatus. Abstract. 41 Introduction. 42 Methods............................................ 45 Results............................... ............. 47 Di scussÎon. 51, Li teta ture Ci ted .................................. 58 lX • CHAPTER 3: Seasonality. environmental lItlccrtajnty and ill!H'ct dOl'l11dIlCV: an empirical model of diapaust" strat0gi0s in Il,,, crickl't Allonemobius fasciatlls. Abstract .............................. id Introduction ..................................
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