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Factors Influencing Mate Choice in Euschistus Conspersus Uhler BEHAVIOR Factors Influencing Mate Choice in Euschistus conspersus Uhler (Heteroptera: Pentatomidae) 1 CHRISTIAN H. KRUPKE, JAY F. BRUNNER, AND VINCENT P. JONES Department of Entomology, Washington State University, Tree Fruit Research and Extension Center, 1100 North Western Ave., Wenatchee, WA 98801 Environ. Entomol. 37(1): 192Ð197 (2008) ABSTRACT The patterns of mating and possible factors inßuencing mate choice in the consperse stink bug, Euschistus conspersus Uhler, were studied in a series of laboratory experiments. Males were found to transfer a signiÞcant percentage of their body mass during the initial mating. Mating was also found to reduce male longevity by 37.8% but had no signiÞcant effect on female longevity or fecundity. There was no evidence of male or female choice based on weight of potential mating partners. There was assortative mating based on experience, however, with males mating preferentially with virgin females and females preferring mated males when given the choice. The implications of theseÞndings within the mating dynamics of this aggregating species are discussed. KEY WORDS Euschistus conspersus, reproductive behavior, mate choice, mating behavior, parental investment The mating period of Euschistus conspersus Uhler has been shown to play a role in the mating dynamics (Heteroptera: Pentatomidae) in northcentral Wash- of stink bugs and a variety of other insect species. For ington occurs once each year, during spring through example, virgin males of the red-shouldered stink bug, early summer. During this time,E. conspersus are often Thyanta pallidovirens Stal, have been shown to donate found in aggregations, typically consisting of 10Ð40 a large fraction (Ϸ17%) of their body mass to the individuals, on host plants (Krupke et al. 2001), a female during the mating sequence (Wang and Millar response that is mediated, at least in part, by a male- 1997), whereas studies of the southern green stink produced aggregation pheromone component (Al- bug, Nezara viridula L., revealed that large males drich et al. 1991, Krupke et al. 2001). However, there mated more often (and with greater fertilization ef- is little published information detailing what actually Þciency) than small males (McLain 1980). In many occurs within these aggregations once they have species of Orthoptera, males make a comparable sac- formed. Mate choice, for example, has not been ex- riÞceÑabout one Þfth of their body weightÑgiven as amined in this species. a spermatophore during mating (Gillette 1904). In To study mate choice in any species, the relative bush-crickets, this spermatophore was found to be roles and contributions of the sexes must Þrst be iden- proportional to the size of the donor, and studies tiÞed. Parental investment theory (Trivers 1972) pre- showed that females showed a preference for larger dicts that females in species where males invest males (Gwynne 1984). In other insect species, masses heavily in offspring Þtness should compete for re- of material have been shown to act as mating plugs that sources and/or for sexually active males and that the enhance mating success by a variety of mechanisms, reverse is true in species where females invest more including inducing behavioral or physiological heavily. In heavy male investment situations, males changes in mated females (Wolfner 1997, Yi Shu and may be expected to exhibit choosiness in mate selec- Gillott 1999), inhibiting or physically blocking further tion. The Þrst step in making predictive hypotheses matings (Monnin and Peeters 1998, Polak et al. 1998), about the roles of males and females in the E. con- and preventing sperm from ßowing out of the female spersus mating system is to document how each sex reproductive tract (Woyciechowski et al. 1994). might maximize its reproductive Þtness. For example, The aforementioned factors may constitute a heavy in male insects of many species, one way of improving male parental investment, perhaps leading male E. Þtness is associated with the mass of material trans- conspersus to exhibit mate choice. To determine ferred to the female during mating. Material transfer whether male mate choice does occur inE. conspersus, we studied one aspect of male parental investment by quantifying the amount of weight change during mat- 1 Corresponding author: Department of Entomology, Purdue University, 901 W. State St., West Lafayette, IN 47907 (e-mail: ing and the effects of mating on male and female [email protected]). longevity. We also describe a series of experiments 0046-225X/08/0192Ð0197$04.00/0 ᭧ 2008 Entomological Society of America February 2008 KRUPKE ET AL.: MATE CHOICE IN E. conspersus 193 examining the possible roles of size and experience in identity of the partner. Times that mating commenced determining male and female mate choice. and ceased were recorded. All insects were weighed immediately before the experiment, and again after mating, or at 0800 hours if they did not mate during the Materials and Methods previous night, and the percentage of weight gained or Weights of Field-Collected Adults.To obtain a mea- lost was calculated for each individual. This protocol sure of the variability within Þeld populations of E. was followed for 6 consecutive d from 10 to 15 May. conspersus, insects were Þeld collected by searching Bugs that were known not to have mated were used as host plants in areas surrounding orchards in Chelan controls. A two-factor ANOVA was used to test for and Douglas counties in Washington state during the interactions between the main factors of sex and mat- period of 25 April through 12 July 2003. Insects were ing status. Differences in percentage weight change sexed and weighed at the WSU Tree Fruit Research with respect to number of matings were analyzed for and Extension Center laboratory in Wenatchee, WA, each sex using single-factor ANOVA. The time mating using a digital scale accurate to 0.1 mg (model 200s; pairs spent in copulae was analyzed using ANOVA as Ohaus, Pine Brook, NJ). Each insect was weighed only well. Percentage weight change data and copulation once, immediately on arrival from the Þeld. The data times were normally distributed and did not require were normally distributed, and weights of Þeld-col- transformation. Where differences were detected lected adults were analyzed using analysis of variance with ANOVA, mean percentage weight changes and (ANOVA) to detect differences between weights of copulation times were analyzed using Fisher least sig- males and females. A two-factor ANOVA was used to niÞcant difference (LSD) test for separation of means test for interaction between the main effects of date (Zar 1999). and weight. Effects of Mating on Longevity and Fecundity. To Male Weight Change During Mating. Adult E. con- determine the effects of mating on longevity in adult spersus were Þeld-collected, as outlined above, from E. conspersus, we placed 30 pairs of virgin, reproduc- areas surrounding orchards in Chelan and Douglas tively mature males and females together in 946-ml counties. Collection began in late March and was containers with mullein leaves as described above. completed by 12 April 2004, immediately after emer- Same-sex pairings of 30 virgin males and 30 virgin gence of the insects from overwintering sites. The females (i.e., 15 pairs of each) were placed in identical insects are relatively inactive during this period, but enclosures during the same period to provide unmated are found basking during warm and sunny days. Dis- controls. These insects had previously been main- sections using criteria developed by Toscano and tained in same-sex greenhouse colonies with mullein Stern (1980) revealed that adult E. conspersus are plants, green beans, and organic sunßower seeds and reproductively immature during this period, and pre- peanuts as food sources. After the outset of the ex- viously collected data show that they typically begin periment, no other food sources (aside from the mul- mating and oviposition in early to mid-May (Alcock lein leaf) were supplied to the insects, to supply water 1971, McGhee 1997). Our early collection periods with only minimal nutrients, thereby simulating an ensured that insects were gathered while they were environment where high-quality food sources (i.e., still virgins. seeds, fruits) were scarce. Insects were held in the Sixty-Þve male and female pairs of E. conspersus same cups for the duration of the experiment, with any were individually placed in 32-oz. transparent plastic dead individuals removed twice weekly. Cups con- cups (Solo Cup Co., Urbana, IL) that had 1-cm-diam- taining male-female pairings were checked nightly eter holes drilled into the cup bottoms. These holes until mating was observed. The total numbers of mat- allowed for the placement of a cut mullein (Verbas- ings over the course of the experiment were not re- cum thapsus L.) leaf into the cup, the stem of which corded. Data collected included the lifespan of each would protrude through the bottom of the cup and insect used during the course of the study and the into a water reservoir below. A mullein leaf was cho- numbers of eggs laid per female per day. This exper- sen because it provided a natural and familiar perch- iment was conducted between 28 May and 13 August ing, mating, and oviposition substrate for the insects 2004. Data were analyzed using two-factor ANOVA to (Krupke et al. 2001). Each cup also contained a single detect interactions between the variables of treatment green bean (Phaesolus vulgaris L.), that was replaced (mating versus same-sex pairs) and sex with respect to twice each week, as well as a small piece of cardboard lifespan. Numbers of eggs laid were analyzed using with several raw organic sunßower (Helianthus ann- single-factor ANOVA to detect differences between uus L) and peanut (Arachis hypogaea L.) seeds glued treatments. Data were normally distributed and did to its surface. Cardboard seed cards were replaced not require transformation.
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