The Cost of Mating: Influences of Life History Traits  and Mating Strategies on Lifespan in Two Closely Related Yponomeuta Species

A. C. Bakker; J. Campos Louçã; P. Roessingh; S. B. J. Menken

doi:10.1155/2011/658908

The Cost of Mating: Influences of Life History Traits and Mating Strategies on Lifespan in Two Closely Related Yponomeuta Species

Bakker, A. C.;Campos Louçã, J.;Roessingh, P.;Menken, S. B. J. 2011-08-09 00:00:00 Hindawi Publishing Corporation International Journal of Zoology Volume 2011, Article ID 658908, 8 pages doi:10.1155/2011/658908 Research Article The Cost of Mating: Inﬂuences of Life History Traits and Mating Strategies on Lifespan in Two Closely Related Yponomeuta Species A. C. Bakker,J.CamposLouc ¸a, ˜ P. Roessingh, and S. B. J. Menken Institute for Biodiversity and Ecosystem Dynamics, University of Amsterdam, P.O. Box 94240, 1090 GE Amsterdam, The Netherlands Correspondence should be addressed to P. Roessingh, roessingh@uva.nl Received 23 December 2010; Revised 21 February 2011; Accepted 18 May 2011 Academic Editor: Marie Herberstein Copyright © 2011 A. C. Bakker et al. This is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. Theory predicts that in monandrous butterﬂy species males should not invest in a long lifespan because receptive females quickly disappear from the mating population. In polyandrous species, however, it pays for males to invest in longevity, which increases the number of mating opportunities and thus reproductive ﬁtness. We tested an extension of this idea and compared male and female lifespan of two closely related Yponomeuta species with diﬀerent degree of polyandry. Our results conﬁrmed the theoretical prediction that male lifespan is ﬁne-tuned to female receptive lifespan; once-mated males and females of both polyandrous species had an equal lifespan. However, the degree of polyandry was not reﬂected in male relative to female lifespan. The observed similar female and male lifespan could largely be attributed to a dramatic reduction of female lifespan after mating. 1. Introduction negative eﬀects, it may seem surprising that multiple mating is the rule in insects at rates that are often much higher Multiple mating can have both positive and negative eﬀects than required for fertilizing the total egg content of a female on female ﬁtness [1–5]. Together these beneﬁts and costs [7]. Therefore, there must also be positive eﬀects associated determine the optimal mating rate of a female. Knowledge with polyandry. These positive eﬀects include compensation of the species-speciﬁc advantages and disadvantages of for mating with a genetically incompatible, inferior, or monandry (females mate only once) and polyandry (females infertile male and an increase in genetic variability among mate more than once) is important for understanding the the oﬀspring ([16, 17], and references therein). Moreover, substances in the male ejaculate (nuptial gifts) can increase adaptive signiﬁcance of mating systems. Negative eﬀects of multiple mating include an increase of energy and time spent female lifespan [18, 19], female fecundity (see [20]for an on mating, as well as the concomitant increased vulnerability overview), and oﬀspring ﬁtness [21, 22]. Positive eﬀects of the ejaculate act either directly, by providing extra resources to predation, sexual diseases, parasites, and pathogens [6]. Furthermore, an excess of sperm can have a negative eﬀect for somatic maintenance of females ([18] and references on both egg production rate (and thus female fecundity) therein [19, 23]), or indirectly, by protecting the female and fertility ([7], and references therein). In addition, in against predators (e.g., the pyrrolizidine alkaloids transferred polyandrous species, mating may reduce female lifespan as by male Utetheisa ornatrix [24]). a consequence of male manipulation [6–9]. Manipulation The mating strategy of a species and multiple mating in comprises both the transfer of toxic compounds, such as particular may inﬂuence certain life history traits such as those found in Drosophila fruitﬂies[10], the bruchid Acan- lifespan. Wiklund et al. [25] suggested that in Lepidoptera diﬀerent mating systems may select for diﬀerent sex-speciﬁc thoscelides obtectus [11], or the nematode Caenorhabditis elegans [12], and physical damage, such as that brought about mortality rates. In species where females mate multiple by bed bugs [13], the bruchid Callosobruchus maculatus times, receptive females are continuously available, and [14], or the dung ﬂy Sepsis cynipsea [15]. Considering these males that allocate resources to increase their lifespan may 2 International Journal of Zoology directly increase their number of opportunities to mate very much alike in many of their life history traits and and consequently their reproductive ﬁtness. Wiklund et al. behaviour. They do however diﬀer in lifespan, and although [25] showed that males of a polyandrous species indeed both are able to mate more than once, they display signiﬁcant lived as long as their conspeciﬁc females while males of diﬀerences in female mating frequencies [39]. The average a monandrous species did not. Males of a monandrous mating frequency of Y. cagnagellus females are 3.0 ± 0.3 times species (under the condition of high synchrony in receptivity (mean ± standard error). In the laboratory, they can mate patterns among individual females) are expected to allocate up to 9 times. The average mating frequency of Y. padellus resources to other traits that improve reproductive success, females is 2.0 ± 0.2, and a majority of females is mating such as development rate, pheromone production, and only once [39]. We can therefore conclude that Y. cagnagellus ﬂight muscles. Fast development leading to males emerging females are highly polyandrous and Y. padellus females have before females (protandry) is advantageous in precopulatory a low degree of polyandry. male competition, whereas male sex pheromones facilitate We ask four questions in this paper: (1) are male and eﬃcient signalling of partners (e.g., in species that form female lifespans equal in polyandrous species as Wiklund male leks [26]) ﬂight muscles improve ﬂight ability and et al. [25] predicts? (2) Can Wiklund’s hypothesis be thus the capability of pursuing and catching females (e.g., in extended to cover sex-speciﬁc adjustment of lifespan related Papilionidae: [27] and in Nymphalidae [28]). to the degree of polyandry of the species? (3) Does mating Here we extend the hypothesis of Wiklund et al. [25]and aﬀect lifespan? (4) Is there a signiﬁcant eﬀect of pupal weight hypothesize that the overall chances of males of encountering on lifespan? a receptive female should aﬀect to which extent resources would be allocated to prolong male reproductive lifespan. 2. Materials and Methods If in a species many females mate only once and few more than once, it is expected that males invest in traits other than In the summer of 2004, ﬁfth instars of the monophagous lifespan (assuming females are receptive around the same Y. cagnagellus were collected in the Netherlands in Cas- ◦ ◦ ◦ ◦ time). We therefore predict that the degree of polyandry will tricum (52 32 N, 4 38 E), Amsterdam (52 8 N, 4 29 E), be reﬂected in the lifespan of males relative to females. ◦ ◦ andOverveen(52 23 N, 4 34 E) from Euonymus europaeus When testing this hypothesis, certain other life history (Celastraceae). Fifth instars of the oligophagous Y. padellus traits and eﬀects of mating need to be considered. First, in ◦ ◦ were collectedinLeiden(52 21 N, 4 56 E) from Crataegus Lepidoptera, positive [19, 29–33], negative [9, 34], or no monogyna (Rosaceae). They were reared as described below eﬀects [35, 36] of mating on lifespan have been observed in on leaves of their host plant. After eclosion, adult moths either sex or in both sexes. Therefore, the eﬀect of mating were placed under a net on their host plant and allowed itself needs to be considered when investigating sex-speciﬁc to mate and lay eggs. Egg batches with ﬁrst instars were longevity and relating it to male investment in lifespan. A sec- kept outdoors under natural conditions. In the next spring, ond aspect that needs to be taken into account is the possible egg batches were opened and the ﬁrst stadium larvae fed eﬀect of body size. In many species, larger individuals tend in Petri dishes (10 cm ∅) with a maximum of 15 L5 to live longer than smaller ones, and females tend to be larger larvae per dish on freshly picked leaves of Prunus spinosa than males ([37] and references therein). Therefore, it may be (Rosaceae) for Y. padellus and E. europaeus for Y. cagnagellus, expected that individuals that have developed from heavier at 22 C, 60% R.H., and L17:D7 photoperiod until pupation. pupae live longer, as they may allocate more resources to the Pupae were weighed individually on an OHAUS Analytical prolongation of adult life. If indeed females are larger than Standard Scale (d = 0.0001 g) microbalance (OHAUS, males, this might cause (all else being equal) lifespan to be Viroﬂay, France) after their pupal skin had hardened. They sex speciﬁc. The inﬂuence of mating itself and possible eﬀects were subsequently placed in individual glass vials (8 cm high of body size on sex-speciﬁc lifespan were unfortunately not and 2 cm ∅) that were closed with a cotton wool plug and investigated by Wiklund et al. [25]. stored in climate boxes at 20 C during the photophase and In this study we reevaluate the hypothesis proposed by 18 C during the scothophase, 70–80% R.H, and L17:D7 Wiklund et al. [25] and speciﬁcally investigate if it holds photoperiod. for species with diﬀerent degrees of polyandry. We used two closely related species from the genus Yponomeuta We used pupal weight as a proxy for body size to test for (Lepidoptera: Yponomeutidae). Yponomeuta cagnagellus a correlation with longevity. For all individuals the number (Hubner) ¨ and Y. padellus (L.) have synchronized female of days between weighing and eclosion was recorded. Pupae eclosion in the ﬁeld and can remate until the end of their were weighed between 3 and 13 days prior to eclosion, with life (A. C. Bakker, unpublished data). Females of both species an average of 6.8 ± 0.1 days for Y. padellus and 8.0 ± 0.2 days mate maximally once per day. Yponomeuta padellus females for Y. cagnagellus. The weight of pupae diminishes every day show synchronized sexual maturity (as indicated by female until eclosion (weight loss amounts to 1.18 ± 0.07% per day ‘calling’ behaviour) and have similar age at ﬁrst mating for Y. cagnagellus and 1.15 ± 0.04% per day for Y. padellus). in the laboratory [38, 39]. Yponomeuta cagnagellus females To correct for this confounding factor, we estimated the also show synchronized sexual maturity [38] but the age of rate of weight loss in the controls of the two species using females mating for the ﬁrst time can vary widely, and this a regression of pupal weight against time to eclosion for may indicate that Y. cagnagellus females are choosy [39]. males and females of both species and used these results to The two species are closely related [40, 41] and therefore predict pupal weight of all insects at the time of eclosion. International Journal of Zoology 3 Table 1: ANOVA table of a GLM model to estimate the eﬀects of sex, species, mating status, and the interaction of sex and species on pupal weight at the time of adult eclosion. Model terms Degree of freedom Deviance Residual degree of freedom Residual deviance FP Null model 502 26345.9 ∗∗∗ Sex 1 6037.5 501 20308.4 207.6 0.0000 ∗∗∗ Species 1 5806.3 500 14502.1 199.7 0.0000 Sex∗species 1 16.5 498 14479.8 0.567 0.4517 Mating status 1 5.8 499 14496.3 0.199 0.6555 ∗∗∗ P< 0.05, P< 0.01, P < 0.001. These estimated weights were subsequently used as a covari- loss since weighing as described above). To test for sex- ate in the GLM analyses to evaluate the eﬀect of pupal speciﬁc lifespan, a sex species interaction term was included. weight (as proxy for adult size) on lifespan. To minimize To test the diﬀerential eﬀects of mating, a sex mating status uncontrolled diﬀerences in weight loss due to variations in and a species∗mating status terms were also included. All humidity, all experiments were conducted in the same room, statistical tests were performed in R (version 2.9.1 [42]). and all treatments were run simultaneously. Throughout the study, average values are presented together After eclosion, moths were sexed and supplied once with their standard errors. a week with a cube of 10% honey-containing 1% agar (w/v; type 1-D LEEO: Sphaero-Q, Leiden, The Netherlands). 3. Results When the moths were sexually mature, that is, at 14–18 days after eclosion for Y. cagnagellus and 8–14 days for Y. 3.1. Pupal Weight. Pupal weights of Y. padellus ranged from padellus, individuals were randomly assigned to either the 17.6 to 48.5 mg for females and from 17.8 to 39.2 mg for mated or the virgin (control) group. One female and one males. In the heavier Y. cagnagellus, pupal weights ranged male were placed together in a Petri dish in the climate room from 28.7 to 57.4 mg for females and for males from 21.1 to at 20-21 C, 60–85% R.H., with a L17 : D7 cycle (of which 46.7 mg. A generalized linear model (Table 1) with estimated the photophase contained 1 h twilight at dawn, provided pupal weight at the day of eclosion as the dependent variable by a 40 W screened lamp). A red darkroom lamp (Philips showed signiﬁcant main eﬀects of both sex and species, PF712E, 15 W) allowed us to observe the moths during but no interaction eﬀect between them, and no eﬀect of the scothophase without disturbing them. The mated group treatment (mated or control). The correlations showed that was allowed to mate once. The females and males of the Y. cagnagellus is signiﬁcantly heavier than Y. padellus and that control group were treated similarly, except that they were females of both species are heavier than males. The lack of placed individually in Petri dishes. Data were collected from amaineﬀect of treatment (mating status) showed that our 43 Y. cagnagellus mating pairs, and 49 unmated males and experimental groups did not systematically diﬀer in weight 51 unmated females served as controls. For Y. padellus, 54 (Figure 1). mating pairs were analysed, and 55 unmated males and 59 unmated females were controls. After mating had ended, 3.2. Lifespan. The individual lifespan of Y. padellus ranged male and female moths were placed individually in glass vials from 15 to 94 days for females (n = 113) and from 19 and were provided with a piece of polystyrene which served to 74 days for males (n = 109). In Y. cagnagellus female as an oviposition substrate. They were kept in a climate lifespan ranged from 17 to 146 days (n = 90) and for males box under the same conditions as when they were maturing from 22 to 115 days (n = 92). We used a generalized linear andwerecontinued to be fedonceaweek.Survivaland model to evaluate the inﬂuence of species, sex, mating status, oviposition were checked daily, with the exception of a few and the covariate pupal weight on lifespan (Table 2). The weekends where checks were made only once in 2 days. model showed signiﬁcant eﬀects for all the factors as well as strong interaction eﬀects (Figure 2, Table 2). Inspection of 2.1. Statistical Analysis. The eﬀects of sex and species on the coeﬃcients for the main eﬀects showed that Y. cagnagellus pupal weight and a possible diﬀerence in pupal weight lived signiﬁcantly longer than Y. padellus, that females lived between treatment groups were investigated using a GLM longer than males, that mating reduced lifespan signiﬁcantly, model that included these factors and a gausian variance and that a higher pupal weight at eclosion increased lifespan. function. The presence of a signiﬁcant interaction of sex and mating To investigate the eﬀects of species, sex, and mating status status revealed however that the eﬀects of mating diﬀered on lifespan, we used a GLM model with a quasipoisson strongly between the sexes. Mating reduced lifespan in both variance function (appropriate for lifespan data that showed sexes but signiﬁcantly more in females than in males. a clear poisson distribution with a tail towards long lifespan). The interaction between sex and species on the other Main eﬀects for species, sex, and mating status were included, hand was not signiﬁcant. While overall the lifespan of Y. as well as the covariate pupal weight (corrected for weight cagnagellus was longer than that of Y. padellus and females 4 International Journal of Zoology Table 2: ANOVA table of a GLM model to estimate the eﬀects of sex, species, mating status, the covariate pupal weight, and speciﬁc planned contrasts. Model term Degree of freedom Deviance Residual degree of freedom Residual deviance FP Null model 502 5055.8 ∗∗∗ Species 1 1381.3 500 3674.5 241.63 0.0000 ∗∗∗ Sex 1 274.0 501 3400.5 47.94 0.0000 ∗∗∗ Mating status 1 273.3 499 3127.2 47.81 0.0000 Pupal weight 1 31.9 498 3095.3 5.58 0.0185 n.s. Sex∗species 1 20.1 497 3075.2 3.52 0.0612 ∗∗∗ Sex∗mating status 1 123.2 496 2952.0 21.55 0.0000 n.s. Species∗mating status 1 6.8 495 2945.2 1.19 0.2765 ∗∗ Sex∗Pupal weight 1 46.7 494 2829.5 8.16 0.0046 ∗ ∗∗ ∗∗∗ P < 0.05, P < 0.01, P < 0.001. 40 80 30 60 20 40 10 20 0 0 Y. cagnagellus Y. padellus Y. cagnagellus Y. padellus Female, control Male, control Female, control Male, control Female, mated Male, mated Female, mated Male, mated Figure 1: Estimated pupal weights (in mg) at the day of eclosion Figure 2: Lifespan (in days) as a function of species (Yponomeuta as a function of species (Yponomeuta padellus or Yponomeuta padellus or Yponomeuta cagnagellus), sex, and mating status (virgin cagnagellus), sex, and mating status (virgin control or once mated). control or once mated). lived longer than males, there was no signiﬁcant sex-speciﬁc 4. Discussion lifespan diﬀerence between Y. padellus (the less polyandrous species) and Y. cagnagellus (the longer living and more 4.1. EﬀectofDegreeofPolyandry on Male Investment polyandrous species). in Lifespan. We investigated sex-speciﬁc lifespan of two Apositivemaineﬀect of pupal weight on lifespan was Yponomeuta species that diﬀer in some life history traits found, but this correlation was also dependent on sex, and and in degree of polyandry. We tested the hypothesis of close examination of the coeﬃcients showed that lifespan was Wiklund et al. [25] that males of polyandrous species invest signiﬁcantly more positively inﬂuenced by pupal weight in in lifespan because this will lead to more matings and thus males than in females. more oﬀspring and higher ﬁtness. We chose species with All the interaction eﬀects described above are also evident adiﬀerent level of polyandry to test if Wiklund’s original in the survival curves presented in Figure 3. The signiﬁcant suggestion would hold even when the degree of polyandry is negative eﬀect of mating on lifespan in the two species considered. Our results conﬁrmed the theoretical prediction is clearly visible, resulting in equal lifespan for males and of polyandrous species having male lifespan close to female females. The interaction eﬀect of mating status with species lifespan: once-mated males and once-mated females had is represented by the relative large reduction of lifespan in an equal lifespan. The hypothesis of Wiklund et al. [25] Y. cagnagellus females after mating. implies that males of polyandrous species continue to live Estimated pupal weight at data of eclosion (mg) Lifespan (days) International Journal of Zoology 5 0.8 0.8 0.6 0.6 0.4 0.4 0.2 0.2 0 20 40 60 80 100 120 140 0 20 40 60 80 100 120 140 Time (days) Time (days) Y. padellus females Y. padellus females Y. cagnagellus females Y. cagnagellus females Y. padellus males Y. padellus males Y. cagnagellus males Y. cagnagellus males (a) (b) Figure 3: (a) Survival curves of virgin adult Yponomeuta padellus and Yponomeuta cagnagellus males and females. Day 0 is the day of eclosion. (b) Survival curves of mated adult Yponomeuta padellus and Yponomeuta cagnagellus males and females. Day 0 is the day of eclosion. after the peak of female receptivity (at least under the higher oviposition rate or to prevent females from remating condition that females receptivity is synchronous, as is the (as is formulated in the adaptive harm hypothesis: [6, 44]). case in Yponomeuta) and thus live past the moment that There are few studies on Lepidoptera in which lifespan of most females have mated once. Indeed, our results showed mated females is compared with virgin female lifespan; in that in both species males live long after the peak of female many studies only lifespan of single-mated females has been receptivity: males of both species live on average more than compared with that of multiple-mated ones. A comparable 45 days after eclosion, but Y. cagnagellus females are receptive negative eﬀect of mating on lifespan was found in Ostrinia at 14.6 ± 1.2 (mean ± SE) days and Y. padellus females at 4.9 nubilalis (Crambidae) [34] and in Colias eurytheme (Pieri- ± 0.6 days after eclosion [39]. dae) [9]. We extended Wiklund’s hypothesis to sex-speciﬁc lifes- While the above explanations picture the reduction in pan diﬀerences between high and low polyandrous species lifespan after mating as a cost of mating, other explanations and expected a larger investment in lifespan in Y. cagnagellus cannot be excluded. The reduction in lifespan could in prin- males; that is, we predicted a signiﬁcant sex by species ciple also be caused by a higher activity of females that were interaction on lifespan. This eﬀect was not found (Table 2) paired up (i.e., activity of females before mating, extensive and while both sex and species inﬂuence lifespan, the eﬀects walking or ﬂying behaviour in search of a suitable oviposition are similar in both species, and we were unable to ﬁnd site after mating). However, in our experimental setup, the evidence for an increase in male investment in lifespan in the moths are placed in Petri dishes and glass vials where they more polyandrous species. cannot easily spend resources on locomotion behaviour. Although we have not formally recorded behaviour, our daily 4.2. Eﬀects of Mating on Lifespan. In Lepidoptera mating can observations showed that moths in all treatments stayed have both positive and negative eﬀects on male and female largely in one place (females call while being motionless and lifespan (e.g., [9, 19, 29–34, 43]). One of our objectives only move around if they do not want to mate), and we have was therefore to investigate if, and how, mating inﬂuenced no reason to believe that paired females were more active lifespan in Y. padellus and Y. cagnagellus.Inour experiment, thanvirginfemales. a single mating event reduced lifespan of both species, but Another explanation for the reduction of lifespan after in particular that of females. Apparently, any advantageous mating could be a change in physiology (i.e., allocation of energy to eggs after fertilisation). Although this would still eﬀects of mating on female lifespan were outweighed by disadvantageous ones. This net negative eﬀect on female represent a cost of reproduction,itwould notbeatrue lifespan might be a side eﬀect (i.e., pleiotropic) of male cost of mating. The design of our experiment is however not suitable to address the diﬀerence between reproduction traits that are beneﬁcial in sperm competition; alternatively, males might deliberately harm females in order to stimulate a and mating. We used an artiﬁcial oviposition substrate, Proportion alive Proportion alive 6 International Journal of Zoology on which only 13 out of 90 Y. cagnagellus and 15 out of 113 in this study, are also expected to inﬂuence sex-speciﬁc Y. padellus females laid eggs (several unmated females also lifespan. Adult feeding can have important eﬀects on sex- oviposited). But even in experiments in which oviposition is speciﬁc mortality rates and sexual behaviour (see also [55]). better facilitated and monitored, it is not easy to discriminate For example, Pararge aegeria (Satyridae) adults lived longer between hypotheses based on physiological changes and when fed [56] and in Pseudoplusia includens (Noctuidae), hypotheses based on harm or male manipulation. Regardless a shortage of food and water led to sex-speciﬁc lifespan of the underlying cause for the eﬀect of mating, we found a in a population in which males normally live as long as signiﬁcant sex∗mating status interaction. A single mating in females [57]. Furthermore, Gotthard et al. [56] found in Yponomeuta did reduce male lifespan signiﬁcantly less than the monandrous P. aegeria adiﬀerence in lifespan between female lifespan even though males of both species transferred males and females when females eclosed synchronously, an ejaculate comprising some 15% of their body weight but no diﬀerence when eclosion of females was continuous during a single mating [45]. Mating also had no eﬀect on throughout the year (causing virgin females to be available male lifespan in Danaus plexippus (Nymphalidae), which over a longer period of time). Sex ratios in the ﬁeld also play transfers an ejaculate that amounts up to 10% of its body an important role because in a population of a monandrous mass [46, 47], nor in Callophrys xami (Lycaenidae), with species with more females than males, it is advantageous for an ejaculate of some 3% of its body mass [48]. However, a male to invest in lifespan as some virgin females will still under resource limitation (achieved by starving the larvae), be available after males have mated once. In monogamous multiple-mated males of C. xami lived shorter than virgin species (in which male lifespan is expected to be shorter than males [48]. We did not test whether resource limitation female lifespan) with protandry (i.e., males eclose or become inﬂuences the eﬀect of mating on lifespan. In our study, as sexually active before females [58]), the earlier eclosion of in that of Oberhauser [47], and in the non-resource limiting males can reduce the diﬀerence in sex-speciﬁc lifespan. This experiment of Cordero [48], adult males were provided with is because male lifespan will tend to be increased to bridge a sugar or honey solution, and this may have enabled them the period between male and female eclosion. To test for to replenish some of their resources. On the other hand, as male investment in lifespan in species with protandry, it is species diﬀer in the allocation of larval and adult resources therefore better to determine if males continue to live after to reproduction and lifespan [49], it is conceivable that in the peak of female receptivity, as we did in this study. Yponomeuta, resources for male lifespan cannot be allocated Our results supported the hypothesis of Wiklund et to reproduction. Another important diﬀerence with the al. [25] that polyandrous males should invest in lifespan studies of Oberhauser [47]and Cordero[48] is that males and therefore live about as long as females. The diﬀer- were mated multiple times, while in this study males were ence in polyandry between the species was however not mated just once. This diﬀerence is important because the reﬂected in male investment in lifespan. The data instead relationship between lifespan and number of matings could strongly suggest a large reduction of female lifespan by be nonlinear. (as yet unknown) negative eﬀects of mating. It would be interesting to elucidate the actual causes, that is, investigate 4.3. Inﬂuence of Pupal Weight on Lifespan. Most Lepidoptera the eﬀects of multiple mating on male and female lifespan acquire most, if not all, of their resources during larval and discriminate between true costs of mating due to male life (so-called capital breeders [50]). These resources can manipulation and male-inﬂicted damage, and changes due in principle be allocated to prolong lifespan, enhance mate to altered resource allocation after mating. ﬁnding, or increase the number and quality of eggs or ejaculate, all of which might increase reproductive output. Acknowledgments By measuring pupal weight, we have indirectly investigated the correlation between resources acquired during the larval The authors thank Louis Lie and Wil van Ginkel for their help stage and adult lifespan, and we expected to ﬁnd a positive with rearing of the Yponomeuta moths and collecting data, correlation between pupal weight and adult lifespan, a and Emiel van Loon for statistical advice. They are grateful to mechanism found across many species ([37] and references Christer Wiklund and two anonymous reviewers for highly therein). 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Copyright: Copyright © 2011 A. C. Bakker et al. This is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.
ISSN: 1687-8477
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DOI: 10.1155/2011/658908
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Abstract

Hindawi Publishing Corporation International Journal of Zoology Volume 2011, Article ID 658908, 8 pages doi:10.1155/2011/658908 Research Article The Cost of Mating: Inﬂuences of Life History Traits and Mating Strategies on Lifespan in Two Closely Related Yponomeuta Species A. C. Bakker,J.CamposLouc ¸a, ˜ P. Roessingh, and S. B. J. Menken Institute for Biodiversity and Ecosystem Dynamics, University of Amsterdam, P.O. Box 94240, 1090 GE Amsterdam, The Netherlands Correspondence should be addressed to P. Roessingh, roessingh@uva.nl Received 23 December 2010; Revised 21 February 2011; Accepted 18 May 2011 Academic Editor: Marie Herberstein Copyright © 2011 A. C. Bakker et al. This is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. Theory predicts that in monandrous butterﬂy species males should not invest in a long lifespan because receptive females quickly disappear from the mating population. In polyandrous species, however, it pays for males to invest in longevity, which increases the number of mating opportunities and thus reproductive ﬁtness. We tested an extension of this idea and compared male and female lifespan of two closely related Yponomeuta species with diﬀerent degree of polyandry. Our results conﬁrmed the theoretical prediction that male lifespan is ﬁne-tuned to female receptive lifespan; once-mated males and females of both polyandrous species had an equal lifespan. However, the degree of polyandry was not reﬂected in male relative to female lifespan. The observed similar female and male lifespan could largely be attributed to a dramatic reduction of female lifespan after mating. 1. Introduction negative eﬀects, it may seem surprising that multiple mating is the rule in insects at rates that are often much higher Multiple mating can have both positive and negative eﬀects than required for fertilizing the total egg content of a female on female ﬁtness [1–5]. Together these beneﬁts and costs [7]. Therefore, there must also be positive eﬀects associated determine the optimal mating rate of a female. Knowledge with polyandry. These positive eﬀects include compensation of the species-speciﬁc advantages and disadvantages of for mating with a genetically incompatible, inferior, or monandry (females mate only once) and polyandry (females infertile male and an increase in genetic variability among mate more than once) is important for understanding the the oﬀspring ([16, 17], and references therein). Moreover, substances in the male ejaculate (nuptial gifts) can increase adaptive signiﬁcance of mating systems. Negative eﬀects of multiple mating include an increase of energy and time spent female lifespan [18, 19], female fecundity (see [20]for an on mating, as well as the concomitant increased vulnerability overview), and oﬀspring ﬁtness [21, 22]. Positive eﬀects of the ejaculate act either directly, by providing extra resources to predation, sexual diseases, parasites, and pathogens [6]. Furthermore, an excess of sperm can have a negative eﬀect for somatic maintenance of females ([18] and references on both egg production rate (and thus female fecundity) therein [19, 23]), or indirectly, by protecting the female and fertility ([7], and references therein). In addition, in against predators (e.g., the pyrrolizidine alkaloids transferred polyandrous species, mating may reduce female lifespan as by male Utetheisa ornatrix [24]). a consequence of male manipulation [6–9]. Manipulation The mating strategy of a species and multiple mating in comprises both the transfer of toxic compounds, such as particular may inﬂuence certain life history traits such as those found in Drosophila fruitﬂies[10], the bruchid Acan- lifespan. Wiklund et al. [25] suggested that in Lepidoptera diﬀerent mating systems may select for diﬀerent sex-speciﬁc thoscelides obtectus [11], or the nematode Caenorhabditis elegans [12], and physical damage, such as that brought about mortality rates. In species where females mate multiple by bed bugs [13], the bruchid Callosobruchus maculatus times, receptive females are continuously available, and [14], or the dung ﬂy Sepsis cynipsea [15]. Considering these males that allocate resources to increase their lifespan may 2 International Journal of Zoology directly increase their number of opportunities to mate very much alike in many of their life history traits and and consequently their reproductive ﬁtness. Wiklund et al. behaviour. They do however diﬀer in lifespan, and although [25] showed that males of a polyandrous species indeed both are able to mate more than once, they display signiﬁcant lived as long as their conspeciﬁc females while males of diﬀerences in female mating frequencies [39]. The average a monandrous species did not. Males of a monandrous mating frequency of Y. cagnagellus females are 3.0 ± 0.3 times species (under the condition of high synchrony in receptivity (mean ± standard error). In the laboratory, they can mate patterns among individual females) are expected to allocate up to 9 times. The average mating frequency of Y. padellus resources to other traits that improve reproductive success, females is 2.0 ± 0.2, and a majority of females is mating such as development rate, pheromone production, and only once [39]. We can therefore conclude that Y. cagnagellus ﬂight muscles. Fast development leading to males emerging females are highly polyandrous and Y. padellus females have before females (protandry) is advantageous in precopulatory a low degree of polyandry. male competition, whereas male sex pheromones facilitate We ask four questions in this paper: (1) are male and eﬃcient signalling of partners (e.g., in species that form female lifespans equal in polyandrous species as Wiklund male leks [26]) ﬂight muscles improve ﬂight ability and et al. [25] predicts? (2) Can Wiklund’s hypothesis be thus the capability of pursuing and catching females (e.g., in extended to cover sex-speciﬁc adjustment of lifespan related Papilionidae: [27] and in Nymphalidae [28]). to the degree of polyandry of the species? (3) Does mating Here we extend the hypothesis of Wiklund et al. [25]and aﬀect lifespan? (4) Is there a signiﬁcant eﬀect of pupal weight hypothesize that the overall chances of males of encountering on lifespan? a receptive female should aﬀect to which extent resources would be allocated to prolong male reproductive lifespan. 2. Materials and Methods If in a species many females mate only once and few more than once, it is expected that males invest in traits other than In the summer of 2004, ﬁfth instars of the monophagous lifespan (assuming females are receptive around the same Y. cagnagellus were collected in the Netherlands in Cas- ◦ ◦ ◦ ◦ time). We therefore predict that the degree of polyandry will tricum (52 32 N, 4 38 E), Amsterdam (52 8 N, 4 29 E), be reﬂected in the lifespan of males relative to females. ◦ ◦ andOverveen(52 23 N, 4 34 E) from Euonymus europaeus When testing this hypothesis, certain other life history (Celastraceae). Fifth instars of the oligophagous Y. padellus traits and eﬀects of mating need to be considered. First, in ◦ ◦ were collectedinLeiden(52 21 N, 4 56 E) from Crataegus Lepidoptera, positive [19, 29–33], negative [9, 34], or no monogyna (Rosaceae). They were reared as described below eﬀects [35, 36] of mating on lifespan have been observed in on leaves of their host plant. After eclosion, adult moths either sex or in both sexes. Therefore, the eﬀect of mating were placed under a net on their host plant and allowed itself needs to be considered when investigating sex-speciﬁc to mate and lay eggs. Egg batches with ﬁrst instars were longevity and relating it to male investment in lifespan. A sec- kept outdoors under natural conditions. In the next spring, ond aspect that needs to be taken into account is the possible egg batches were opened and the ﬁrst stadium larvae fed eﬀect of body size. In many species, larger individuals tend in Petri dishes (10 cm ∅) with a maximum of 15 L5 to live longer than smaller ones, and females tend to be larger larvae per dish on freshly picked leaves of Prunus spinosa than males ([37] and references therein). Therefore, it may be (Rosaceae) for Y. padellus and E. europaeus for Y. cagnagellus, expected that individuals that have developed from heavier at 22 C, 60% R.H., and L17:D7 photoperiod until pupation. pupae live longer, as they may allocate more resources to the Pupae were weighed individually on an OHAUS Analytical prolongation of adult life. If indeed females are larger than Standard Scale (d = 0.0001 g) microbalance (OHAUS, males, this might cause (all else being equal) lifespan to be Viroﬂay, France) after their pupal skin had hardened. They sex speciﬁc. The inﬂuence of mating itself and possible eﬀects were subsequently placed in individual glass vials (8 cm high of body size on sex-speciﬁc lifespan were unfortunately not and 2 cm ∅) that were closed with a cotton wool plug and investigated by Wiklund et al. [25]. stored in climate boxes at 20 C during the photophase and In this study we reevaluate the hypothesis proposed by 18 C during the scothophase, 70–80% R.H, and L17:D7 Wiklund et al. [25] and speciﬁcally investigate if it holds photoperiod. for species with diﬀerent degrees of polyandry. We used two closely related species from the genus Yponomeuta We used pupal weight as a proxy for body size to test for (Lepidoptera: Yponomeutidae). Yponomeuta cagnagellus a correlation with longevity. For all individuals the number (Hubner) ¨ and Y. padellus (L.) have synchronized female of days between weighing and eclosion was recorded. Pupae eclosion in the ﬁeld and can remate until the end of their were weighed between 3 and 13 days prior to eclosion, with life (A. C. Bakker, unpublished data). Females of both species an average of 6.8 ± 0.1 days for Y. padellus and 8.0 ± 0.2 days mate maximally once per day. Yponomeuta padellus females for Y. cagnagellus. The weight of pupae diminishes every day show synchronized sexual maturity (as indicated by female until eclosion (weight loss amounts to 1.18 ± 0.07% per day ‘calling’ behaviour) and have similar age at ﬁrst mating for Y. cagnagellus and 1.15 ± 0.04% per day for Y. padellus). in the laboratory [38, 39]. Yponomeuta cagnagellus females To correct for this confounding factor, we estimated the also show synchronized sexual maturity [38] but the age of rate of weight loss in the controls of the two species using females mating for the ﬁrst time can vary widely, and this a regression of pupal weight against time to eclosion for may indicate that Y. cagnagellus females are choosy [39]. males and females of both species and used these results to The two species are closely related [40, 41] and therefore predict pupal weight of all insects at the time of eclosion. International Journal of Zoology 3 Table 1: ANOVA table of a GLM model to estimate the eﬀects of sex, species, mating status, and the interaction of sex and species on pupal weight at the time of adult eclosion. Model terms Degree of freedom Deviance Residual degree of freedom Residual deviance FP Null model 502 26345.9 ∗∗∗ Sex 1 6037.5 501 20308.4 207.6 0.0000 ∗∗∗ Species 1 5806.3 500 14502.1 199.7 0.0000 Sex∗species 1 16.5 498 14479.8 0.567 0.4517 Mating status 1 5.8 499 14496.3 0.199 0.6555 ∗∗∗ P< 0.05, P< 0.01, P < 0.001. These estimated weights were subsequently used as a covari- loss since weighing as described above). To test for sex- ate in the GLM analyses to evaluate the eﬀect of pupal speciﬁc lifespan, a sex species interaction term was included. weight (as proxy for adult size) on lifespan. To minimize To test the diﬀerential eﬀects of mating, a sex mating status uncontrolled diﬀerences in weight loss due to variations in and a species∗mating status terms were also included. All humidity, all experiments were conducted in the same room, statistical tests were performed in R (version 2.9.1 [42]). and all treatments were run simultaneously. Throughout the study, average values are presented together After eclosion, moths were sexed and supplied once with their standard errors. a week with a cube of 10% honey-containing 1% agar (w/v; type 1-D LEEO: Sphaero-Q, Leiden, The Netherlands). 3. Results When the moths were sexually mature, that is, at 14–18 days after eclosion for Y. cagnagellus and 8–14 days for Y. 3.1. Pupal Weight. Pupal weights of Y. padellus ranged from padellus, individuals were randomly assigned to either the 17.6 to 48.5 mg for females and from 17.8 to 39.2 mg for mated or the virgin (control) group. One female and one males. In the heavier Y. cagnagellus, pupal weights ranged male were placed together in a Petri dish in the climate room from 28.7 to 57.4 mg for females and for males from 21.1 to at 20-21 C, 60–85% R.H., with a L17 : D7 cycle (of which 46.7 mg. A generalized linear model (Table 1) with estimated the photophase contained 1 h twilight at dawn, provided pupal weight at the day of eclosion as the dependent variable by a 40 W screened lamp). A red darkroom lamp (Philips showed signiﬁcant main eﬀects of both sex and species, PF712E, 15 W) allowed us to observe the moths during but no interaction eﬀect between them, and no eﬀect of the scothophase without disturbing them. The mated group treatment (mated or control). The correlations showed that was allowed to mate once. The females and males of the Y. cagnagellus is signiﬁcantly heavier than Y. padellus and that control group were treated similarly, except that they were females of both species are heavier than males. The lack of placed individually in Petri dishes. Data were collected from amaineﬀect of treatment (mating status) showed that our 43 Y. cagnagellus mating pairs, and 49 unmated males and experimental groups did not systematically diﬀer in weight 51 unmated females served as controls. For Y. padellus, 54 (Figure 1). mating pairs were analysed, and 55 unmated males and 59 unmated females were controls. After mating had ended, 3.2. Lifespan. The individual lifespan of Y. padellus ranged male and female moths were placed individually in glass vials from 15 to 94 days for females (n = 113) and from 19 and were provided with a piece of polystyrene which served to 74 days for males (n = 109). In Y. cagnagellus female as an oviposition substrate. They were kept in a climate lifespan ranged from 17 to 146 days (n = 90) and for males box under the same conditions as when they were maturing from 22 to 115 days (n = 92). We used a generalized linear andwerecontinued to be fedonceaweek.Survivaland model to evaluate the inﬂuence of species, sex, mating status, oviposition were checked daily, with the exception of a few and the covariate pupal weight on lifespan (Table 2). The weekends where checks were made only once in 2 days. model showed signiﬁcant eﬀects for all the factors as well as strong interaction eﬀects (Figure 2, Table 2). Inspection of 2.1. Statistical Analysis. The eﬀects of sex and species on the coeﬃcients for the main eﬀects showed that Y. cagnagellus pupal weight and a possible diﬀerence in pupal weight lived signiﬁcantly longer than Y. padellus, that females lived between treatment groups were investigated using a GLM longer than males, that mating reduced lifespan signiﬁcantly, model that included these factors and a gausian variance and that a higher pupal weight at eclosion increased lifespan. function. The presence of a signiﬁcant interaction of sex and mating To investigate the eﬀects of species, sex, and mating status status revealed however that the eﬀects of mating diﬀered on lifespan, we used a GLM model with a quasipoisson strongly between the sexes. Mating reduced lifespan in both variance function (appropriate for lifespan data that showed sexes but signiﬁcantly more in females than in males. a clear poisson distribution with a tail towards long lifespan). The interaction between sex and species on the other Main eﬀects for species, sex, and mating status were included, hand was not signiﬁcant. While overall the lifespan of Y. as well as the covariate pupal weight (corrected for weight cagnagellus was longer than that of Y. padellus and females 4 International Journal of Zoology Table 2: ANOVA table of a GLM model to estimate the eﬀects of sex, species, mating status, the covariate pupal weight, and speciﬁc planned contrasts. Model term Degree of freedom Deviance Residual degree of freedom Residual deviance FP Null model 502 5055.8 ∗∗∗ Species 1 1381.3 500 3674.5 241.63 0.0000 ∗∗∗ Sex 1 274.0 501 3400.5 47.94 0.0000 ∗∗∗ Mating status 1 273.3 499 3127.2 47.81 0.0000 Pupal weight 1 31.9 498 3095.3 5.58 0.0185 n.s. Sex∗species 1 20.1 497 3075.2 3.52 0.0612 ∗∗∗ Sex∗mating status 1 123.2 496 2952.0 21.55 0.0000 n.s. Species∗mating status 1 6.8 495 2945.2 1.19 0.2765 ∗∗ Sex∗Pupal weight 1 46.7 494 2829.5 8.16 0.0046 ∗ ∗∗ ∗∗∗ P < 0.05, P < 0.01, P < 0.001. 40 80 30 60 20 40 10 20 0 0 Y. cagnagellus Y. padellus Y. cagnagellus Y. padellus Female, control Male, control Female, control Male, control Female, mated Male, mated Female, mated Male, mated Figure 1: Estimated pupal weights (in mg) at the day of eclosion Figure 2: Lifespan (in days) as a function of species (Yponomeuta as a function of species (Yponomeuta padellus or Yponomeuta padellus or Yponomeuta cagnagellus), sex, and mating status (virgin cagnagellus), sex, and mating status (virgin control or once mated). control or once mated). lived longer than males, there was no signiﬁcant sex-speciﬁc 4. Discussion lifespan diﬀerence between Y. padellus (the less polyandrous species) and Y. cagnagellus (the longer living and more 4.1. EﬀectofDegreeofPolyandry on Male Investment polyandrous species). in Lifespan. We investigated sex-speciﬁc lifespan of two Apositivemaineﬀect of pupal weight on lifespan was Yponomeuta species that diﬀer in some life history traits found, but this correlation was also dependent on sex, and and in degree of polyandry. We tested the hypothesis of close examination of the coeﬃcients showed that lifespan was Wiklund et al. [25] that males of polyandrous species invest signiﬁcantly more positively inﬂuenced by pupal weight in in lifespan because this will lead to more matings and thus males than in females. more oﬀspring and higher ﬁtness. We chose species with All the interaction eﬀects described above are also evident adiﬀerent level of polyandry to test if Wiklund’s original in the survival curves presented in Figure 3. The signiﬁcant suggestion would hold even when the degree of polyandry is negative eﬀect of mating on lifespan in the two species considered. Our results conﬁrmed the theoretical prediction is clearly visible, resulting in equal lifespan for males and of polyandrous species having male lifespan close to female females. The interaction eﬀect of mating status with species lifespan: once-mated males and once-mated females had is represented by the relative large reduction of lifespan in an equal lifespan. The hypothesis of Wiklund et al. [25] Y. cagnagellus females after mating. implies that males of polyandrous species continue to live Estimated pupal weight at data of eclosion (mg) Lifespan (days) International Journal of Zoology 5 0.8 0.8 0.6 0.6 0.4 0.4 0.2 0.2 0 20 40 60 80 100 120 140 0 20 40 60 80 100 120 140 Time (days) Time (days) Y. padellus females Y. padellus females Y. cagnagellus females Y. cagnagellus females Y. padellus males Y. padellus males Y. cagnagellus males Y. cagnagellus males (a) (b) Figure 3: (a) Survival curves of virgin adult Yponomeuta padellus and Yponomeuta cagnagellus males and females. Day 0 is the day of eclosion. (b) Survival curves of mated adult Yponomeuta padellus and Yponomeuta cagnagellus males and females. Day 0 is the day of eclosion. after the peak of female receptivity (at least under the higher oviposition rate or to prevent females from remating condition that females receptivity is synchronous, as is the (as is formulated in the adaptive harm hypothesis: [6, 44]). case in Yponomeuta) and thus live past the moment that There are few studies on Lepidoptera in which lifespan of most females have mated once. Indeed, our results showed mated females is compared with virgin female lifespan; in that in both species males live long after the peak of female many studies only lifespan of single-mated females has been receptivity: males of both species live on average more than compared with that of multiple-mated ones. A comparable 45 days after eclosion, but Y. cagnagellus females are receptive negative eﬀect of mating on lifespan was found in Ostrinia at 14.6 ± 1.2 (mean ± SE) days and Y. padellus females at 4.9 nubilalis (Crambidae) [34] and in Colias eurytheme (Pieri- ± 0.6 days after eclosion [39]. dae) [9]. We extended Wiklund’s hypothesis to sex-speciﬁc lifes- While the above explanations picture the reduction in pan diﬀerences between high and low polyandrous species lifespan after mating as a cost of mating, other explanations and expected a larger investment in lifespan in Y. cagnagellus cannot be excluded. The reduction in lifespan could in prin- males; that is, we predicted a signiﬁcant sex by species ciple also be caused by a higher activity of females that were interaction on lifespan. This eﬀect was not found (Table 2) paired up (i.e., activity of females before mating, extensive and while both sex and species inﬂuence lifespan, the eﬀects walking or ﬂying behaviour in search of a suitable oviposition are similar in both species, and we were unable to ﬁnd site after mating). However, in our experimental setup, the evidence for an increase in male investment in lifespan in the moths are placed in Petri dishes and glass vials where they more polyandrous species. cannot easily spend resources on locomotion behaviour. Although we have not formally recorded behaviour, our daily 4.2. Eﬀects of Mating on Lifespan. In Lepidoptera mating can observations showed that moths in all treatments stayed have both positive and negative eﬀects on male and female largely in one place (females call while being motionless and lifespan (e.g., [9, 19, 29–34, 43]). One of our objectives only move around if they do not want to mate), and we have was therefore to investigate if, and how, mating inﬂuenced no reason to believe that paired females were more active lifespan in Y. padellus and Y. cagnagellus.Inour experiment, thanvirginfemales. a single mating event reduced lifespan of both species, but Another explanation for the reduction of lifespan after in particular that of females. Apparently, any advantageous mating could be a change in physiology (i.e., allocation of energy to eggs after fertilisation). Although this would still eﬀects of mating on female lifespan were outweighed by disadvantageous ones. This net negative eﬀect on female represent a cost of reproduction,itwould notbeatrue lifespan might be a side eﬀect (i.e., pleiotropic) of male cost of mating. The design of our experiment is however not suitable to address the diﬀerence between reproduction traits that are beneﬁcial in sperm competition; alternatively, males might deliberately harm females in order to stimulate a and mating. We used an artiﬁcial oviposition substrate, Proportion alive Proportion alive 6 International Journal of Zoology on which only 13 out of 90 Y. cagnagellus and 15 out of 113 in this study, are also expected to inﬂuence sex-speciﬁc Y. padellus females laid eggs (several unmated females also lifespan. Adult feeding can have important eﬀects on sex- oviposited). But even in experiments in which oviposition is speciﬁc mortality rates and sexual behaviour (see also [55]). better facilitated and monitored, it is not easy to discriminate For example, Pararge aegeria (Satyridae) adults lived longer between hypotheses based on physiological changes and when fed [56] and in Pseudoplusia includens (Noctuidae), hypotheses based on harm or male manipulation. Regardless a shortage of food and water led to sex-speciﬁc lifespan of the underlying cause for the eﬀect of mating, we found a in a population in which males normally live as long as signiﬁcant sex∗mating status interaction. A single mating in females [57]. Furthermore, Gotthard et al. [56] found in Yponomeuta did reduce male lifespan signiﬁcantly less than the monandrous P. aegeria adiﬀerence in lifespan between female lifespan even though males of both species transferred males and females when females eclosed synchronously, an ejaculate comprising some 15% of their body weight but no diﬀerence when eclosion of females was continuous during a single mating [45]. Mating also had no eﬀect on throughout the year (causing virgin females to be available male lifespan in Danaus plexippus (Nymphalidae), which over a longer period of time). Sex ratios in the ﬁeld also play transfers an ejaculate that amounts up to 10% of its body an important role because in a population of a monandrous mass [46, 47], nor in Callophrys xami (Lycaenidae), with species with more females than males, it is advantageous for an ejaculate of some 3% of its body mass [48]. However, a male to invest in lifespan as some virgin females will still under resource limitation (achieved by starving the larvae), be available after males have mated once. In monogamous multiple-mated males of C. xami lived shorter than virgin species (in which male lifespan is expected to be shorter than males [48]. We did not test whether resource limitation female lifespan) with protandry (i.e., males eclose or become inﬂuences the eﬀect of mating on lifespan. In our study, as sexually active before females [58]), the earlier eclosion of in that of Oberhauser [47], and in the non-resource limiting males can reduce the diﬀerence in sex-speciﬁc lifespan. This experiment of Cordero [48], adult males were provided with is because male lifespan will tend to be increased to bridge a sugar or honey solution, and this may have enabled them the period between male and female eclosion. To test for to replenish some of their resources. On the other hand, as male investment in lifespan in species with protandry, it is species diﬀer in the allocation of larval and adult resources therefore better to determine if males continue to live after to reproduction and lifespan [49], it is conceivable that in the peak of female receptivity, as we did in this study. Yponomeuta, resources for male lifespan cannot be allocated Our results supported the hypothesis of Wiklund et to reproduction. Another important diﬀerence with the al. [25] that polyandrous males should invest in lifespan studies of Oberhauser [47]and Cordero[48] is that males and therefore live about as long as females. The diﬀer- were mated multiple times, while in this study males were ence in polyandry between the species was however not mated just once. This diﬀerence is important because the reﬂected in male investment in lifespan. The data instead relationship between lifespan and number of matings could strongly suggest a large reduction of female lifespan by be nonlinear. (as yet unknown) negative eﬀects of mating. It would be interesting to elucidate the actual causes, that is, investigate 4.3. Inﬂuence of Pupal Weight on Lifespan. Most Lepidoptera the eﬀects of multiple mating on male and female lifespan acquire most, if not all, of their resources during larval and discriminate between true costs of mating due to male life (so-called capital breeders [50]). These resources can manipulation and male-inﬂicted damage, and changes due in principle be allocated to prolong lifespan, enhance mate to altered resource allocation after mating. ﬁnding, or increase the number and quality of eggs or ejaculate, all of which might increase reproductive output. Acknowledgments By measuring pupal weight, we have indirectly investigated the correlation between resources acquired during the larval The authors thank Louis Lie and Wil van Ginkel for their help stage and adult lifespan, and we expected to ﬁnd a positive with rearing of the Yponomeuta moths and collecting data, correlation between pupal weight and adult lifespan, a and Emiel van Loon for statistical advice. They are grateful to mechanism found across many species ([37] and references Christer Wiklund and two anonymous reviewers for highly therein). 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The Cost of Mating: Influences of Life History Traits and Mating Strategies on Lifespan in Two Closely Related Yponomeuta Species

The Cost of Mating: Influences of Life History Traits and Mating Strategies on Lifespan in Two Closely Related Yponomeuta Species

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The Cost of Mating: Influences of Life History Traits and Mating Strategies on Lifespan in Two Closely Related Yponomeuta Species

The Cost of Mating: Influences of Life History Traits and Mating Strategies on Lifespan in Two Closely Related Yponomeuta Species

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