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Survivorship and Growth of Sexually and Asexually Derived Larvae of Alsophila Pometaria (Lepidoptera: Geometridae)

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Survivorship and Growth of Sexually and Asexually Derived Larvae of Alsophila Pometaria (Lepidoptera: Geometridae) University of Nebraska - Lincoln DigitalCommons@University of Nebraska - Lincoln Faculty Publications in the Biological Sciences Papers in the Biological Sciences 6-1985 Survivorship and Growth of Sexually and Asexually Derived Larvae of Alsophila pometaria (Lepidoptera: Geometridae) Lawrence G. Harshman University of Nebraska - Lincoln, [email protected] Douglas J. Futuyma State University of New York Follow this and additional works at: https://digitalcommons.unl.edu/bioscifacpub Part of the Developmental Biology Commons, and the Entomology Commons Harshman, Lawrence G. and Futuyma, Douglas J., "Survivorship and Growth of Sexually and Asexually Derived Larvae of Alsophila pometaria (Lepidoptera: Geometridae)" (1985). Faculty Publications in the Biological Sciences. 273. https://digitalcommons.unl.edu/bioscifacpub/273 This Article is brought to you for free and open access by the Papers in the Biological Sciences at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Faculty Publications in the Biological Sciences by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. Harshman & Futuyma in American Naturalist (June 1985) 125(6). Copyright 1985, University of Chicago. Used by permission. SURVIVORSHIP AND GROWTH OF SEXUALLY AND ASEXUALLY DERIVED LARVAE OF ALSOPHILA POMETARIA (LEPIDOPTERA: GEOMETRIDAE) A substantialbody of theoryis devoted to understandingthe relative advan- tages of sexual and asexual reproduction.It is generallyunderstood that asexual formspotentially have a higherrate of reproductionbecause theysave the cost of producing males. The microevolutionaryconsequences of sexual and asexual reproductionare less clear. Sexual reproductiongenerates abundant genotypic diversitywhich may be adaptivelyadvantageous (Williams 1975; Maynard Smith 1978). Asexual reproductionmay perpetuate combinations of genes that are coadapted (Templeton 1979), heterotic(Suomalainen et al. 1976; White 1979), or specialized (Vrijenhoek1979, 1984). Thus, it is possible thatthe fitnessof a sexual populationmay be lower thanan asexual, in part,because recombinationtends to break up especially favorable genotypes (Williams 1975; Hebert 1978). If it is generallyobserved thatasexual reproductionhas an immediateadaptive as well as a reproductiveadvantage, then it is difficultto see how sexual reproductioncan be maintainedby short-termadvantages (Williams 1975). A comparisonof closely relatedsexual and asexual formsis a promisingavenue of researchto evaluate experimentallythe consequences of both modes of repro- duction (Maynard Smith 1978). In this study,geometrid moth larvae (Alsophila pometaria) derivedfrom both kindsof reproductionwere rearedon differenthost plants. The goal was to assess larval viabilityand growthin an ecologically relevantcontext and thus partiallycharacterize the fitnessof sexual and asexual reproduction. Biological Background Alsophila pometaria, the sole Nearctic memberof its genus (Inoue 1943), is a species of moth whose unusual mode of reproductionhas been studied exten- sively withelectrophoretic markers. Most females are gynogeneticand requirea matingwith a nonspecificmale in orderto reproduce.Gynogenetic females of the fall cankerwormtransmit the maternalgenotype intact to theirprogeny (Mitter and Futuyma 1977). A varietyof electrophoreticallydefined multilocus genotypes (clones) are foundin asexual populations,and it has been establishedthat there is a coexistingsexual population(Mitter et al. 1979). Electrophoreticsurveys have revealed thatthe same alleles are presentat similarfrequencies in the sexual and asexual populations (Mitteret al. 1979; Harshman 1982). The species is univoltine.Eggs are laid on treesmostly in earlywinter and hatch about the time of budbreak in the spring.There can be extensive dispersal of hatchlinglarvae by ballooningon silk threads(Futuyma et al. 1981). This larval Am. Nat. 1985. Vol. 125, pp. 8%-902. ? 1985 by The Universityof Chicago. 0003-0147/85/2506-0010$02.00.All rightsreserved. NOTES AND COMMENTS 897 dispersalcommonly may resultin movementamong different host trees(Futuyma et al. 1981). The larvae feed on the foliage of numerousdeciduous tree species untilearly Junewhen they drop to the groundand pupate. The correspondence between the maturationschedule of the larvae and that of the foliage on which theyfeed is criticalbecause, like the European wintermoth Operophtera brumata (Feeny 1970), theyare unable to completegrowth on excessively maturefoliage. MATERIALS AND METHODS Identificationof ReproductiveMode Larvae forthese experimentswere obtainedfrom egg masses foundon trees or directlyfrom mating adults collected in the field.The collectionsites were wood- lots dominatedby oak (Quercus spp.) and red maple (Acer rubrum)near the Stony Brook campus of the State Universityof New York, which is located on the northernshore of Long Island. A numberof eggs fromeach egg mass were set aside at room temperaturefor early hatchingand subsequent electrophoretic analysis. The remainderwere leftoutside for the winterin 1-dramglass vials to be used in experimentsafter larvae hatched in the spring. It is not possible to determinethe reproductivemode of adultfemales from their externalmorphology. Thus, it is our procedureto electrophoresea numberof sibs froman egg mass to determinedirectly the mode of reproductionof the mother. First-instarlarvae that hatched in the lab were homogenizedin 12-14 kl of the grindingbuffer used by Mitteret al. (1979). Afterelectrophoresis, we stainedfor two of the polymorphicenzymes employed for adults (PGM and PGI). Usually, 10 larvae were run from each egg mass. Sexual reproductionwas detected by observingrecombination of electrophoreticmarkers among larval sibs (Harshman 1982). Larvae of specific asexual lineages were isolated fromfemales whose multilocusgenotype correspondedto that of an established clone (Mitteret al. 1979; Futuymaet al. 1981). Growthin a ControlledEnvironment In the spring of 1980, larvae from a sexually derived sibship (HL75) were compared to two asexual lineages. One lineage was the widespreadand abundant asexual genotype2(B), which is associated withoak in the fieldand is apparently an oak specialist (Mitteret al. 1979). The second was asexual genotype48(Z), which is abundantin mixed oak-and-maplestands. Mitteret al. (1979) suggested thatthis genotypemay be more generalizedin its feedinghabits than the oak- or maple-associated clones. Larvae of the maple-specialistgenotype, 26(A), were not available for this experiment. Larvae were rearedentirely on eitheroak or red maple leaves obtainedfrom the field.All larvae reared on oak were exposed to a mixtureof two species (Quercus coccinea and Q. velutina). Hatchlinglarvae fromeach sibship employed in this study were split evenly between oak and maple foliage. The experimentwas conductedwith approximately 10 larvae per 5-litercardboard container. Food was 898 THE AMERICAN NATURALIST replenishedevery otherday in the formof freshsprigs of foliage,with the twigs insertedinto flasks of water. The containerswere placed in an environmental chamber adjusted for 18 h of lightat 24? C and 6 h of darkness at 16? C. Larval survivorshipwas not recorded in the experimentbecause a major source of mortalitywas uncontrolleddesiccation of first-instarlarvae. After3 wk to 1 mo of growth,prepupal larvae leftthe foliageand began burrowing.When an individual reached this stage its weightwas recorded. Prepupal weightis correlatedwith fecundityin thefall cankerworm (Schneider 1979),and generallypre-image weight is highlycorrelated with fecundity in insects (Engelmann 1970). Body size is not necessarilya primarydeterminant of fitness,but it is correlatedwith fecundity, which is a componentof fitness. Survivorshipand Growthof Larvae in the Field In the springof 1981 an experimentwas conductedto evaluate the survivorship and growthof sexually and asexually derived larvae. The tested genetic groups consisted of a sexual sibship (VW1 18), the oak-specialist [2(B)] and maple- specialist [26(A)] genotypes,and two clones thatare uncommonon Long Island. The site of the studywas a woodlot on the Stony Brook campus in which five oak trees (Quercus coccinea) and five red maple trees were selected as host plants. Soon afterbudbreak, larvae thathad just hatchedwere placed on foliage enclosed by finemesh bags as describedby Futuymaet al. (1981). Each host tree supportedat least two bags withthree larvae fromeach geneticgroup. Afterapproximately 1 mo of growth,when larvae began to reach the prepupal stage, all rearingbags were broughtin fromthe field,and larval survivorshipwas recorded. Larvae thathad not attainedthe prepupal stage were raised for a few days in the laboratoryon vegetationsimilar to thatavailable to themin the field. All prepupal larvae were allowed to pupate in a mixtureof sterilizedsand and pottingsoil. After2 mo, pupae were removed fromtheir subterranean cocoons and sexed. They were thenlyophilized and the dryweight of each individualwas determined. Several aspects of these studiesdeserve comment.First, they are not designed to quantifytotal fitness.Moore (1976), in a comparisonof the gynogeneticfish Poeciliopsis monacha-occidentalis with a related bisexual species, describes three aspects of fitness.One is the twofoldreproductive advantage accruing to asexual reproduction,another is mating success, and the third is "primary fitness,"which includes fecundity and survival.In the presentstudy on Alsophila pometaria, we are concerned with measures that
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