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Head-Capsule Widths and Number of Larval Instars in Gynaepiwra Species ...38 Vi National tibrary Bibliothèque nationale du Canada Acquisitions and Acqui=i?ionset Bibliographic Services seivices bibliographiques 395 Wellingtort Street 393. rue Wellington OttawaON KiAON4 OttawaON K1AON4 canada Canada The author has granted a non- L'auteur a accordé une licence non exclusive licence allowing the exclusive permettant a la National Library of Canada to Bibliothèque nationale du Canada de reproduce, loan, distribute or sell reproduire, prêter, distribuer ou copies of this thesis in microfom, vendre des copies de cette thèse sous paper or electronic formats. la forme de rnicrofiche/film, de reproduction sur papier ou sur format électronique. The author retains ownership of the L'auteur conserve la propriété du copyright in this thesis. Neither the droit d'auteur qui protège cette thèse. thesis nor substantiai extracts fiom it Ni la thèse ni des extraits substantiels may be printed or otherwise de celle-ci ne doivent être imprimés reproduced without the author's ou autrement reproduits sans son permission. autorisation. Supervisor: Dr. Richard A. Ring Increases in temperature and precipitation predicted under global wann i ng are expected to be most pronounced and thus have their greatest impact on ecosystems at high latitudes. Insects constitute a major component of the foodwebs of terrestrial ecosystems and should be among the Tirst organisms to show noticeable responses to predicted global warming, especiaily in the Arctic where climatic conditions are often limiting. However, interactions among species must also be taken into account. The genus GynaepIwra Hübner (Lepidoptera: Lymantnidae) is represented in North Amenca by two species, G. groenhdica (Wocke) and G. rossii (Curtis), and their geographic distributions overlap broadly across the Canadian Arctic. Previous studies have examined the biology, ecology, and physiology of these two species and have revealed many adaptations to the Arctic environment, but the immature stages of these insects have been misidentified even in recently published reports. Both species are found at Alexandra Fiord, Ellesmere Island, a High Arctic oasis largely isolated by expanses of ocean and icecap, and the population of G. groeiihrzdica at this site is thought to be limited mainly by prasitoid-induced mortality nther than by climatic conditions. Field observations, surveys, and tempenture-manipulation expenments were conducted at Alexandra Fiord during the spring and summer of 1994, 1995, and 1996; Iaboratory reaing was conducted under controlled conditions at the University of Victoria in the spring of 1996 and 1997. Immature stages of both species of Gyriaepiwra were described and illustrated, and ail species of insect parasitoids using GymzepIwra species as hosts at Alexandra Fiord were identified. Life histones and seasonal phenologies for Gyri~epIwruspecies and their insect parasitoids were elucidated from field studies, and temperanire/development reiationships for selected stages of most of these species were denved from Iaboratory rearing. The results of field studies and laboratoiy rearing were compared and used to formulate predictions about the responses of these insects to predicted global warming. Immature stages of the twc species of Gynaeplwra are easily distinguished by differences in the colour patterns, fom, and overall length of the larval hairs and by the structure of their cocoons. Both species of Gyr~aeplwracomplete metamorphosis and reproduction within a single growing season but spread larval development over a number of years. In G. groenlardica, seven larval instars and annual moulting combilis to produce a seven year lire cycle whereas G. rossii develops through six larval instars at a rate of two or three moulu per year, resulting in a three or four year life cycle. The parasitoid complex at Alexandra Fiord consists of three primas, parasitoids, Hyposoter pectirzatus (Thomson) (Hymenoptera: Ichneumonidae), ExoristB nsp. (Dipten: Tachinidae), and Ciierogetza gelida (Coquillett) (Diptera: Tachinidae), and one hyperpansitoid, Crypti~~leechi Mason (Hymenoptera: Ichneumonidae). Al1 of the parasi toids are uni vol tine, al thoug h H. pectinatirs may undergo delayed develo pment in some cases, and each of the primary parasitoids relies primarily on a single larval instar for hosts w hereas the hyperparasi toid attacks the primary parasi toi& dunng thei r metamorphosis. Seasonal phenologies of the parasitoids provide optimal access to new hosts but parasitoid-avoidance stmtegies of Gynaepiiora larvae ensure ihat a proportion of their populations escape pansitism. Laboratory rearing showed that the relative timing of host and parasitoid seasonal phenologies is maintained over a broad range of temperatures; therefore, temperature inmeases p~dictedunder global warming are unlikely to have any great effect on host-parasitoid interactions. However, increased cloudiness associateci with the predicted increase in precipitation might have profound effects resulting from lower ground-level temperatures caused by a Iack of solar heating. The extent of this effect is uncertain but might lead to reproductive fidure in Gynaephora species, with similar repercussions for the insect pamsitoids. TABLE OF CONTENTS TlTLE PAGE .................................. i ABSTRACT .................................. ü TABLE OF CONTENTS ............................ v LIST OF TABLES ..............................."ii LISTOFFIGURES .............................. xi ACKNOWLEDGEMENTS ........................... .!a DEDICATION ................................."i INTRODUCTION ............................... 1 The Greenhouse Effect andOlobal Warming Scenarios .............. 1 Implications of Global Wanning for Insects .................... 4 Global Warming and Insects in the Arctic ..................... 6 The Experirnental Animals ............................ 9 North Amencan Gynaepiwra Species ..................... 9 Insect Parasitoids of North Amencan Gynaeplwra Species ........... 13 The Study Site ................................. 18 Objectives ...................................30 MATERIALS AND METHODS .........................21 Field Studies ..................................71 Field Surveys ................................24 Field Evperiments .............................. 26 Gymeplwra life histories and effect of temperature on development .....26 Reproductive isolation of Gy~ephorospecies ...............31 Efrect of temperature on primary parasitoid metamorphosis .........31 Voltinism of the prirnary parasitoids ....................33 Laboratory Rearing ...............................35 Head-capsule Widths and Number of Larval Instars in Gynaepiwra Species ...38 vi RESULTS ................................... 42 Descriptions of Immafurc Stases of Gymeplwra Species .............. 42 Eggs ....................................42 Larvae ................................... 42 Hibernacula ................................. 50 Cocoons .................................. 53 Pupae .................................... 5Q Life Histories and Biology ............................ 56 GytlaeplwraSpecies ............................. 58 Head-capsule widths and number of larval instars .............. 58 Larval activity and rnoulting frequency ................... 65 Metamorphosis and reproduction ..................... 73 Parasi toids of Gyrzaepjwra S pecies ...................... 77 Hyposoter pectit1atu.s (Hymenoptem Ichneumonidae) ............ 81 fioriskz n.sp. (Diptera: Tachinidae) .................... 86 Cf~togeirngelida (Dipten: Tachiniche) ................... 93 Cryptzcs leeclu (Hymenoptera: Ichneumonidae) ............... 99 Temperature/Development Relationshijzi ....................103 FieId Studies ................................103 iaboratory Rearing .............................109 IXSCUSSION ................................. 121 Identification cf Immature Stages of Gymzepiwra Species .............121 Life Histories and Biology ..........................124 Gyrzaepiwra Species ...........................124 Head-capsule widths and number of larval instars .............124 Larval activity and moulting frequency ...................127 Metamorphosis and reproduction .................133 Parasi toi& of GynaepfwraSpecies .....................136 Hyposoter pectinnfirs (Hym enopten: Ic hneumonidae) ............139 Erorisa n.sp. and Cf~togerzagelida (Diptem Tachinidae) ..........142 Cryptirs kecfti (Hymenoptera: Ichneumonidae) .............145 Temperature/Developrnent Relationships ..................147 Field Studies ................................147 Labontory Rearing .............................149 Life History Strategies ..............................154 Implications of GloM Warming .........................161 CONCLUSIONS ................................164 REFERENCES CITED .............................167 APPENDIX: Instar Determination From Measured Head-capsule Widths . 187 LIST OF TABLES Table 1. 1nsect parasitoids reported to use North A mencan Gynaeplwra s pecies as hosts. Table 2. Morphological differences berneen hi& arctic Gytiueplwra species in the immature stages. For measurements, the full range found in this study is given. Table 3. Cornparison of previousl y ~portedmean larval head capsule widths (HCW)of Gyrraeplwra groetdarrdica to expected and observed mean HCW determined in this study, with corresponding growth ratios and boundary points between observed means. Reported means are from Kukal and Kevan (L987), expected means were predicted by the Brooks-Dyar Rule, observed means
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