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The UniversÍty of Manitoba BIONOMICS 0F THE SUNFLOWER BEETLE, Zygognamma exclamationis (F. ) (COLfOptsRA:CHRYSOMELIDAE), AND ITS PAMSITES IN MANTTOBA by Gannet Bruce Neill A Thesis Submitted to the Facutty of Gnaduate Studies In Par"tiat Fulfillment of the Requinements for the Degnee of Docton of Philosophy Depantment of Entomology Winnipeg, ManÍtoba T982 BIONOMICS OF TIIE SUNFLOI'IER BEETLE, ZYGOGRAI"ÍMA EXCLA},IATIONIS (F. ) (COLEOPTERA:CHRYSOMELIADAE) , AND ITS PARASITES IN MANITOBA BY GARNET BRUCE NEILL A thesis submitted to the Faculty of Graduate Studies of the University of Manitoba irr partial fulfillment of the requirements of the degree of DOCTOR OF PHILOSOPHY o 1982 Permissiort has been granted to the LIBRARY OF THE UNIVER- SITY OF MANITOBA to lend or sell copies of this thesis, to the NATIONAL LIBRARY OF CANADA to microfilm this thesis a¡ld to lend or sell copies of the filnr, and UNIVERSITY MICROFILMS to publish an abstract of this thesis. The author reserves other publication rights, and neither the thesis nor extensivc extracts from it may be prirrted or other- wise reproduced without the author's written permission. 11 ABSTRACT The sunflowen beetle, Zygognarnrna exclamationis (F.), has one genenation per yean in Manitoba. The beetle ovenwintens as a sexually immatr::re adul-t in soil, usually at a depth of 15 cm or less, in fiel-ds in which it completed lanval development. Mating and egg-laying do not take plaee príon to hibennation. The mean date of finst emengence of beetles, ovell a foun-yean pe::iod, was May 20, the nange being May 11 to 27. In one year, ovenwintening beetles emerged oven a peniod of 24 days with peak emengence on day 13. A method fon pr.edíetíng the time of emengence of ovenwintering beetles was developed using a system of thenmat unit (fU) accumul-ations based on soil- temperature necordings fuom a depth of 5 cm. Males emenged one to foun days before females duning the initial emergence peniod, but no diffenences wene noted thereaften. Mal-es and fenales wene distinguishable by the shape of the fifth abdominal ster"nite. Sexual matur"ation was initiated eithen befone o:r sho::t1y afte:: emergence fuom hibernation. Posthibennation sexual development was napid, mating being noted within two days of finst emelrgence and egg- laying within one week of eme::gence. BehavÍoun duning cor:r'tship was simple and basic, and both males and females mated frequentty thnoughout the oviposition peniod. Orce oviposition was initiated, females laid eggs almost every ,, day during the oviposition peniod. Eggs wene laid on the leaves and stem of the host plant and wene usually found singly on ín ir"negular iii groups. FemaLes neared in the labor"atony at 25oC laid an avenage of 30.7 eggs per" day drring an ovíposition pe:riod of 60.7 days. Egg pnoduction pen female avenaged 11965 with a nange of 260 to 31587. In field tnials, the average daily egg pnoduction was 15.2, oven a mean oviposition peniod of 47.1 days. The mean egg pnoduction in two field tnials was 719 with a nange of !32 to 1,554. Eggs wene pnesent ín commencial sunfl-owe:: fields fon six to seven weeks. Most beetles died shortly aften oviposition ceased. The íncubation peniod of the eggsr which are clream to yellow and measu::e 1.33 by 0.61 mm, vras affected by tempenature above 9.goC, the lowen th¡eshold tempenatune for development (ToLO); the estinated the::nal r:nit nequirement fon incubation was 83. Lar"vae passed thnough four instans distinguishable by width of the head capsule. As with incr:batíon, tempenatuï,e sÍgníficantly affected the dur"ation of the lanval stage. The estimated ToLO fon the finst- to fou::th-instans was 8.1, B.9o 7.5 and 3.8oC, nespectively, and estimated TU nequinement at the corresponding ToLOts was 76, 50, 63 and 83, respectively. A ToLO of 8.4oC râras necortrnended for pnedictíng lanval development in the field. Over. a thnee-yean peniod, the mean date of first and last appearance of larvae in seeded sunflowen fields was Jur.e 10 and July 28 fon a mean dr:ration of 51 days. The mid-point in the development of the second-instar was detenmined as the optimal time fon contnol of Larvae sirrce at that time most eggs had hatched and líttte foliage damage had occunned. Methods lv of accunately pnedicting this poínt, based on field obsenvations and TU accumulations wene developed. Following completion of the fou::th instan, larvae entened the soil to pupate. The duration of the pnepupal and pupal periods combined was 16.9 days at 20oC, with an estimated ToLO of 7.4oC and TU nequir"ement of 2tt. The total developmental time fnom egg to adult was 44.6 days at 20oC. Upon emengence fuom pupation, p::ehíbernation adults fed fon one to three weeks pnion to entry into the soil to ovenwinter". The mean date of finst appearance of pnehibernation adults was 'JuIy 26, 49 days or" 555 TUfs (7.4oC ToLO) aften the finst lanvae were obsenved. Pnehibennation adults did not fly, nate on lay eggs. A numben of diffe::ent predators attacked the differ"ent life stages of the sunflowen beetle. HippodamÍa tnedecimpunctata tibialis (Say) and Collops vittalus Say fed on eggs, Hippodamia convergens Guenin-Menevil-le and g¡r:ysopa. caranea Stephens fed on eggs and lanvaeo Lebia atniventnis Say, Perilliodes bioculatus (Fab.) and Nabis sp. fed on lanvae, and Podisus maculiventnis (Say¡ and the blackbir"d, Agelaius phoeniceus phoeniceug L. fed on adults. Fou:r' species of panasites vrene neared fuorn diffenent stages in the life cycle of the sunflower beetle; Myiopharus sp. fnom adults, Enixestus winnemana Cr"awford f::on eggs, and Doryphonophaga macella Reinhand and D. donyphonae (Riley) from larvae. v Myiophanus sp.: ê tachinid reaned fnom adults, has two genenations per year on the sunflowen beetle. The panasite ovenwintens as a finst-Ínstan naggot in the abdomen of the ovenwinteníng host, migrating to the bnain in spning and eventually killing the host shontly before the time of normal emellgence. The parasite emel:ges about one month after. nonmal emergence of the hosto in time to panasitize the last of the posthibernation beetles to for"m the second genenation whÍch in tunn develops in time to attack pnehibennation beetles in late su¡nmerl and eanly fa1I. The maggots deposíted by second genenation adults form the ovenwintening population. In fÍeld collections of pnehibennation beetles oven thnee years, incidence of panasitism varied from 0.2 to 17.1 pencent. 0n1y one panasite was neared fnom each beetle. The hypenpanasiteo Pe::ilarnpus sp., þras ::eaned from pupanía of Myiophanus sp. A one-month delay in spning emengence of the panasite followÍng that of the beetl-e pnovídes an oppor"tunity fon cont::ol of posthibernation beetles with nininal effect on Myiophanus sp. Ênixestus winnemanara pteromalid, was reaned fnom eggs of the sunflower beetle. The life cycle of this panasite was completed in 16.1 days at 20oC, so sever.al genenations perî yean are possible. Howeven, only one or possibly two genenations ane completed in any given sunflowen field because of the timited peniod during which host eggs are available. Panasitism by E. winnenana nangeC up to 16.8 pencent in commercial sunflowen fields, while up to 65.7 pencent panasitism was vi obsenved in small plots used fon an oviposition tnial fon the sunflowen beetle. Adults of E. winnemana are present ín the field at the same time as are adult;r ar" beetle. Contr"o1 of the host with """rrowen insecticides would. thus pose a hazand for the panasite. Donyphonophaga macella, a tachinid, panasitized la¡vae in all stages by subcutaneous lanviposition. Appeanance of the ovenwintened genenation is well synchnonized with that of the host, par"asitisrn oceunning within a week of first appearance of host lanvae. Females of D. macell-a ar?e highly feeund, depositing a mean of 284.6 maggots over a !4.2 day lanviposition peniod in the labonato::y. Supenpanasitism was common, but only one panasite successfully developed in each host 1arva. The par.asite passes th::ough three instans, distinguishable by the size and shape of the nouth-hooks, the naggot r"enaining as a fÍr"st instan until the host larva completes deveLopment. The panasite develops napidly aften the host lan¡a enters the soil to pupate, kiIlíng the host in two on thnee days and emenging as a ner^r-genenation ad.utt about the same time as pnehibernation beetles. This generation is poor"ly synchronized wíth the host since the population of host lanvae would be declining napídly at the tíme of emergence of the new genenation of par.asites fncidence of panasitism in the field, over a five-year peniod, r:anged fnom 0.1 to oven 70 pencent, the majonity of the maggots being deposited du:ring the eanly lanva1 peniod when finst- and second-instan lanvae vrerle prevalent. D. macella tended to act in an inverse v11 density-dependent mannerâ; as the density of the host lar:vae. increased, the nelatíve impact of D. macella decr:eased. The hypenpanasite, Penilanpus sp. was neaned from pupania of D. macella. Control of host lanvae with insecticides would effectively el-ininate the panasite in any tneated. anea because D. macella initiates lanviposition veny soon aften first emergence of host lanvae. D. donyphonae, a tachinid, was of rníno:: impontance as a panasíte of the sunflowe¡'beetle, nepresenting onJ-y 0.2 pencent of the panasites neared fnom host lanvae. Par"asites and pnedators are impo::tant natr:na1 control agents of the sunflowen beetle in Manítoba, and aid in suppnessing pest populations ín most yeans.