Menken, S.B.J.

Parker, K.

Roessingh, P.

English

Many male Lepidoptera transfer a large ejaculate, containing nutrients and/or secondary plant compounds. In principle this 'nuptial gift' could help to increase female survival and longevity. Here we investigated the effects of both multiple mating and extra nutrition in the adult stage on males and females in two species of Yponomeuta. Mated females had significantly shorter live spans than unmated females and similar fecundity. Females therefore do not appear to benefit from remating, which suggests that the considerable weight transfer of the males to the females during mating does not provide nutritional value. Females of Y. cagnagellus, which received honey throughout their adult lifetime, lived significantly longer and laid larger egg batches than females, that did not receive honey. In Y. padellus this effect was not found. It is concluded that neither species benefits from multiple mating and that in Yponomeuta cagnagellus adult nutrition rather then mating increases female reproductive output and longevity, which contrasts with many studies of Macrolepidoptera

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UvA-DARE is a service provided by the library of the University of Amsterdam (https://dare.uva.nl)UvA-DARE (Digital Academic Repository)Effects of multiple mating and adult nutrition on longevity and fecundity in twoYponomeuta speciesParker, K.; Roessingh, P.; Menken, S.B.J.Publication date2008Document VersionFinal published versionPublished inProceedings of the Netherlands Entomological Society MeetingLink to publicationCitation for published version (APA):Parker, K., Roessingh, P., & Menken, S. B. J. (2008). Effects of multiple mating and adultnutrition on longevity and fecundity in two Yponomeuta species. Proceedings of theNetherlands Entomological Society Meeting, 19, 115-120. http://www.nev.nl/sete/sete-2008/115-120-Parker-2008.pdfGeneral rightsIt is not permitted to download or to forward/distribute the text or part of it without the consent of the author(s)and/or copyright holder(s), other than for strictly personal, individual use, unless the work is under an opencontent license (like Creative Commons).Disclaimer/Complaints regulationsIf you believe that digital publication of certain material infringes any of your rights or (privacy) interests, pleaselet the Library know, stating your reasons. In case of a legitimate complaint, the Library will make the materialinaccessible and/or remove it from the website. Please Ask the Library: https://uba.uva.nl/en/contact, or a letterto: Library of the University of Amsterdam, Secretariat, Singel 425, 1012 WP Amsterdam, The Netherlands. Youwill be contacted as soon as possible.Download date:09 Mar 2023Katherine Parker, Peter Roessingh & Steph B.J. MenkenUniversity of Amsterdam, Institute for Biodiversity and Ecosystem Dynamics, POBox 94062, 1090 GB Amsterdam, The Netherlands, E-mail: roessingh@uva.nlMany male Lepidoptera transfer a large ejaculate, containing nutrientsand/or secondary plant compounds. In principle this ‘nuptial gift’could help to increase female survival and longevity. Here we investi-gated the effects of both multiple mating and extra nutrition in theadult stage on males and females in two species of Yponomeuta. Matedfemales had significantly shorter live spans than unmated females andsimilar fecundity. Females therefore do not appear to benefit from re-mating, which suggests that the considerable weight transfer of themales to the females during mating does not provide nutritional value.Females of Y. cagnagellus, which received honey throughout their adultlifetime, lived significantly longer and laid larger egg batches thanfemales, that did not receive honey. In Y. padellus this effect was notfound. It is concluded that neither species benefits from multiple mat-ing and that in Yponomeuta cagnagellus adult nutrition rather then mat-ing increases female reproductive output and longevity, which con-trasts with many studies of Macrolepidoptera.Keywords: nuptial gift, honey, reproductive output, MacrolepidopteraFemale butterflies and moths require significant amounts of energy forpheromone production, calling, courtship, and oviposition. Many empiricalstudies in Macrolepidoptera have found that, in addition to adult nutrition, mul-tiple mating can have a positive effect on female longevity and fecundity. A sin-gle mating can provide a female with enough sperm to fertilize all her eggs(Lederhouse 1981). Remating is therefore not needed for fertilisation. However,males can transfer an ejaculate to the female that contains a ‘nuptial gift’, acces-sory substances, which provide nourishment to the female, increasing her fecun-dity (Cook & Wedell 1996).In addition to direct nutritional effects that increase fecundity, nuptial giftsmight also increase female lifespan. This benefits the female, presuming that thelonger a female lives the more opportunities she has to produce offspring and findsuitable partners that can provide a genetic benefit to her offspring in the form ofEffects of multiple mating and adult nutritionon longevity and fecundity in two YponomeutaspeciesPROC. NETH. ENTOMOL. SOC. MEET. - VOLUME 19 - 2008 115116genetic variation, bet hedging for imperfect mate assessment, and higher levels ofsperm competition (Tores-Vila & Walker 1980, Halliday 1983, Watson 1991).Most studies look at the reproductive behavior of butterflies, few havefocused on moths. An exception is the study by De Jong (1988) who found insmall ermine moths that females who do not receive honey before mating hadstunted growth, small egg number, produce less attractive pheromones, and yetwere more successful in mating repeatedly. We therefore designed an experi-ment in Yponomeuta moths with the following questions in mind:(1) How do multiple mating and adult nutrition affect female longevity, mat-ing frequency, and egg production in two closely related species of Yponomeuta?(2) Can nutritionally stressed females increase their longevity and reproduc-tive output by mating more frequently?Both species feed on their own specific host-plant: Yponomueta padellus can befound feeding on plants from various species of Rosaceae including Prunus spin-osa while Yponomeuta cagnagellus exclusively feeds on Euonymus europaeus belong-ing to the Celastraceae (Menken 1992). In late spring females deposit up to 150eggs in clusters near a bud or side branch of their food plant and eggs hatch threeweeks later (Povel 1984).We tested the potential benefits of nuptial gifts on fecundity and longevityby mating females with one or two males or by providing continuous access tomales. In addition, the effect of honey was investigated. We quantified the totalnumber of matings, the insect’s total lifespan, and the total surface of all eggbatches laid over a female’s complete lifetime.MATERIAL AND METHODSMating and feeding treatmentsA total of 389 Y. cagnagellus and 396 Y. padellus males and females were assignedto unmated (‘virgin’), once-mated, twice-mated, and unlimited mating groups.Half of the insects were fed honey throughout their lifetime and half were not.Both mating and courtship were recorded once per hour between 9:00 and18:00 hours. A mating was recorded when the male and female remained in cop-ula for more than 1.5 hours. Females that did not want to mate were suppliedwith a new virgin male twice per week.Egg production analysis & longevityMost studies examining the importance of male investment for female repro-duction assume that the mass of the male ejaculate is positively correlated withfemale output. Spermatophore size is however not the best measurement of maleinvestment, since it is the content and not the size of the spermatophore thatdetermines its true value (Marshall 1989, Delisle & Bouchard 1995). For this rea-son we measured lifetime female fecundity rather than spermatophore size. Ahost-plant stem was placed with every female in a Petri-dish and was checkedEVOLUTIONARY ECOLOGY3-5 times per week for egg batches. Batches were photographed, and the eggbatch surface was measured using the program ‘ImageJ’ (Rasband 1997).Longevity was measured from the moment a moth enclosed till its death for allmoths in both experiments.RESULTSAdult NutritionThe lifespan for Y. cagnagellus was approximately one month and for Y. padellusit was approximately two months. Yponomeuta cagnagellus females did utilizehoney to increase their lifespan (Fig. 1; ANOVA: df=1, F=15.3, p<0.001). Honeydid not increase the lifespan of the shorter lived Y. padellus (Fig. 2; ANOVA:df=1, F=1.82, p=0.18).Yponomeuta cagnagellus females with access to honey laid significantly moreegg batches and also produced a significant larger egg batch surface (Fig. 3;K. PARKER, P. ROESSINGH & S.B.J. MENKEN117Figure 1. Effect of mating frequency onlongevity of Yponomeuta cagnagellus forunmated (n=61), once-mated (n=47),twice-mated (n=43), and free-mating (3+)females (n=48). Approximately half thenumber of females assigned to the fourmating groups (0-3+) received honeywhilst the other half did not.Figure 2. Effect of mating frequency onlongevity in Yponomeuta padellus forunmated (n=60), once-mated (n=49),twice-mated (n=38), and free-mating (3+)females (n=49). Approximately half thenumber of females assigned to the fourmating groups (0-3+) received honeywhilst the other half did not.ANOVA: df=1, F=7.78, p<0.01) than females without this extra source of energy.Yponomeuta padellus females on the other hand did not benefit significantly fromtheir access to honey and both groups produced about equal egg batch surfaces(Fig. 4; ANOVA: df=1, F=0.14, p=0.91).Multiple MatingMultiple mating did not increase female longevity in either species ofYponomeuta. Mating not only failed to extend female lifespan but in fact signif-icantly reduced it (Figs. 1 and 2; ANOVA, df=1, Fcag=17.49 and Fpad=43.05,P<0.001).Total egg production (both egg batch number and batch surface) did notchange significantly when females were mated multiple times (Figs. 5 and 6;ANOVA, df=1, Fcag=0.08 and Fpad=3.13, P=ns for both species). Unmated femalesof both species in all cases laid only very few (unfertilized) eggs.DISCUSSIONOur results show that contrary to expectation (based on literature data forMacrolepidoptera), multiple mating has a negative effect on longevity andfemales did not convert the content of the male spermatophore into eggs.EVOLUTIONARY ECOLOGY118Figure 3. Effect of honey on total eggbatch surface for mated Yponomeutacagnagellus females that received (n=26) ordid not receive honey (n=22) and for vir-gin females that received (n=31) or did notreceive honey (n=30).Figure 4. Effect of honey on total eggbatch surface for mated Yponomeuta padel-lus females that received (n=24) or did notreceive honey (n=25) and for virginfemales that received (n=30) or did notreceive honey (n=30).Lederhouse et. al (1990) observed that Papilio glaucus ingest nectar, sodium, andother electrolytes and transfer this extra source of nutrition to females duringmating in order to increase female reproductive output. Arnqvist & Nilsson(2000) suggested that multiple mating may increase female fitness by stimulat-ing egg production, replenishing sperm supplies, and offering accessory sub-stances that contribute to female survival and/ or reproduction. Here we foundno support for these ideas as mating multiply had a significant negative effecton female longevity and did not affect reproductive output. It should be notedhowever that the lack of effect of the nuptial gift as an energy source does notrule out that it might contain secondary plant compounds that increase the sur-vival of the female and/or the offspring.In contrast to the lack of a nutritional benefit from multiple mating, honeycan be used as an energy source by Y. cagnagellus to extend lifespan and producelarger eggs batches. Yponomeuta padellus on the other hand did not profit fromhoney. This is possibly due to time constraints related to its shorter lifespan. Itseems that Y. cagnagellus has the ability to use external carbohydrate sources inits adult life, while Y. padellus females are relying exclusively on their larvalresources.K. PARKER, P. ROESSINGH & S.B.J. MENKEN119Figure 5. Effect of mating on total eggbatch surface produced by Yponomeutacagnagellus for unmated (n=61), once-mated (n=47), twice-mated (n=43), andfree-mating(3+) females (n=48).Approximately half the number offemales assigned to the four matinggroups (0-3+) received honey whilst theother half did not.Figure 6. Effect of mating on total eggbatch surface produced by in Yponomeutapadellus for unmated (N=60), once-mated(N=49), twice-mated (N=38), and free-mating (3+) females (N=49).Approximately half the number offemales assigned to the four matinggroups (0-3+) received honey whilst theother half did not.REFERENCESArnqvist G. & Nilsson T. 2000. The evolution of polyandry: multiple-mating andfemale fitness in insects. Animal Behavior 43:559-567.Cook, P. & Wedell, N. 1996. Ejaculate dynamics in butterflies: a strategy for maximiz-ing fertilization success? Proceedings of the Royal Scociety Londen B 263: 1047-1051.De Jong, M.C.M. 1988. Evolutionary approaches to insect communication system. PhDthesis, pp.163. Leiden Univ. Leiden.Delisle, J. & Bouchard, A. 1995. Male larval nutrition in Choristoneura rosaceana(Lepidoptera: Tortricidae): an important factor in reproductive success. Oecologia104:508-517.Halliday, T.R. 1983. The study of mate choice. In Male Choice (ed. P. Bateson), pp 32.Cambridge University Press.Lederhouse, R.C. 1981. The effect of mating frequency on egg fertility in the black swal-lowtail, Papilio polyxenes asterius (Papilionidae). Journal of the Lepidopterists’ Society35:266-277.Lederhouse, R.C. & Ayres, M.P. & Scriber, J.M. 1990. Adult nutrition affects male viril-ity in Papilio glaucus. L. Functional Ecology 4:743-751.Marshall, L.D. & McNeil, J.N. 1989. Spermatophore mass as an estimate of male nutri-ent investment: a closer look in Pseudaletia unipuncta (Hayworth)(Lepidoptera:Noctuidae). Functional Ecology. 3:605-612.Menken, S.B.J. 1992. Small ermine moths (Yponomeuta): Their host relations and evolu-tion. Annual Review of Entomology 37:41-66.Povel, G.D.E. The identification of the European small ermine moths, with special refe-rence to the Yponomeuta padellus-complex (Lepidoptera, Yponomeutidae).Proceedings of the Koninklijke. Nedederlamdse Akademie van. Wetenschappen Serie C87:149-80.Rasband, W.S. ImageJ. U. S. National Institutes of Health, Bethesda, Maryland, USA,http://rsb.info.nih.gov/ij/, 1997-2006.Tores-Vila L. & Walker, W.F. 1980. Sperm utilization strategies in nonsocial insects.American Naturalist 115:780-799.Watson, P.J. 1991. Multiple paternity as genetic bet-hedging in female sierra dome spi-ders (Linyphia litigiosa: Linyphiidae). Animal Behavior 41:135-148.EVOLUTIONARY ECOLOGY120

Effects of multiple mating and adult nutrition on longevity and fecundity in two Yponomeuta species

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Effects of multiple mating and adult nutrition on longevity and fecundity in two Yponomeuta species

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