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Relationship to Mate Fecundity in the Southern Green Stinkbug, Nezaraviridula (Hemiptera: Pentatomidae)

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Relationship to Mate Fecundity in the Southern Green Stinkbug, Nezaraviridula (Hemiptera: Pentatomidae) Heredity 64 (1990) 161—167 The Genetical Society of Great Britain Received 25 July 1989 Male copulatory success: heritability and relationship to mate fecundity in the southern green stinkbug, Nezaraviridula (Hemiptera: Pentatomidae) Denson Kelly MCL&n* Department of Biology, Landrum Box 8042, Nancy Brannen Marsh Georgia Southern College, Statesboro, Georgia 30460, U.S.A. Single male southerngreen stinkbugs, Nezaraviridula, sequestered in mating chambers with six females varied in the number of copulations achieved. Females copulating with relatively successful males were more fecund than females copulating with less successful males. This suggests that females mating successful males receive greater nutritional rewards per mating or that their risk of contagion is reduced. There was a significant correlation between the mating success of fathers and sons. Since egg quality (hatch rate) and egg size (diameter) did not covary with male copulatory success, this correlation indicates additive genetic variation for male mating. success in a natural population. Thus, for females there appears to be a positive relationship between offspring quality and mate effects on fecundity. Additive genetic variation for mating success may be maintained by countervailing mortality selection on males imposed by a parasitoid which is attracted by male pheromones. INTRODUCTION tiveness on female fecundity and offspring repro- ductive successareunknown for any species. The Sexualselectionarises fromvariationinthe num- present study examines both the heritability of ber ofmates among individualsof the same sex male mating success and the correlation between (Darwin,1871). Female choice contributesto vari- male success and female fecundity in the southern ation in the mating success of males in many organ- green stinkbug, Nezaraviridula (L.).Thus,the isms (Trivers, 1985), including some insects present study addresses whether or not female (Thornhill and Alcock, 1983). Female preferences choice for genes promoting offspring competitive- are selected to maximize immediate fecundity ness is necessarily at odds with choice for male (Kirkpatrick, 1985) and may entail choice for investment (e.g., Burley, 1988; Weatherhead and males offering nutritional rewards that defray the Robertson, 1979). cost of egg production (e.g., Butlin et a!., 1987; Sakaluk, 1985; Gwynne, et a!., 1984). Females also may be selected to choose males on the basis of METHODS their potential genetic contributions to progeny Male copulatory success (Hamilton and Zuk, 1982; Searcy, 1982; Borgia, 1979). Females possessing and exercising preferen- Southern green stinkbugs, N. viridula, were reared ces for attractive mates might have more attractive in the laboratory from the eggs of 30 females sons than nonchoosy females (Heisler, 1985). collected in mid-April (1988) from wild mustard Additive genetic variation forattractivemale growing in an abandoned pasture in Clito, Georgia traitsexists in some insect populations (Hedrick, (U.S.A.). Forty sets of six virgin females and one 1988 and references therein). This genetic variation virgin male, 10 days old as adults at the initiation orcorrelatedgenetic variation maybe expressed of the experiment, were housed in 1-litre mason inthe competitiveness (Partridge, 1980; Taylor et jars. Copulating pairs were sequested into -litre a!.,1987)and, ultimately, reproductive success of cartons where the mated female remained for the offspring. However, the dual effects of male attrac- duration of her life. The mated male was returned to the unmated females remaining in the jar from *To whomcorrespondence should be addressed. which he came. Both jars and cartons were 162 D. K. McLAIN AND N. B. MARSH supplied with two pole beans which served as both Male copulatory success could be inherited a food and water source. Fresh beans were sub- nongenetically as a consequence of nutrient trans- stituted for old beans every 5 days. Insects were fer from male to female during copulation which maintained on a 14L: IOD photophase at 20°C and impacts egg size or quality. Larger eggs might give 80 per cent RH. Males and females were placed rise to larger more competitive progeny (e.g., in jars at random with regard to size. All seven Johnson,1982). Therefore, the percentage of eggs adults housed together had different mothers. This hatching and egg diameter were determined as, arrangement reduced the potential for inbreeding respectively, indices of egg quality and size. Egg depression among progeny used in the heritability diameter was measured with an ocular micrometer estimates described below. Since experimental (1 unit=001 mm) at the widest point across the females did not share the same mother, they had operculum, after hatching. Five eggs from the first been reared in different jars. Thus, any inadvertent egg mass laid by the first female mated were environmental influence during development on measured, two from the middle of one of the outer the willingness of females to mate should have had egg rows, two from the middle of the innermost minimal effect on the number of females a male egg row, and one positioned between the other mated. Since the placement of jars on racks in the two sets. environmental chamber was randomized daily and Variation in male reproductive success can since all jars were supplied identically, variation derive from the fecundity of females mated as well in the number of females mated was not due to as the number mated (Wade and Arnold, 1980). environmental variation. Relatively attractive or vigorous males may be Male-male competition could not be expressed more successful at obtaining copulations with in the present experiment. Therefore, differences more fecund females. Alternatively, relatively among males in the number of females mated may unattractive males may be selected to devote more be ascribed to variation in the attractiveness and time and energy to the courtship of more fecund staying power or general vigour of males. Copula- females. Given these possibilities and given that tory success of a male is measured as the number females do vary tremendously in egg productivity of females mated within 30 days after the initiation (unpublished observations), the heritability of of the experiment. Males copulating with all six male reproductive success was also assessed. females within 30 days were supplied with a Reproductive success is defined as the total number seventh female. However, none of the males mated of eggs laid by all the females a male mated. a seventh time. In addition to the number of Heritability was estimated from the regression of females copulated, the size of each male was also son reproductive success on father reproductive determined. Here, male size refers to the width success. Variation in reproductive success poten- across the pronotum at the level of the lateral tially reflects contributions from variation in male angles. Width was measured with an ocular attractiveness and vigour, female fecundity and micrometer to the nearest 0125 mm. Measure- longevity, and offspring fitness (reflecting parental ments were made after the experiment had con- investment and genetic constitution). Reproduc- cluded. tive success was determined for 22 father/son pairs. Heritabilityof male success Female Theheritability of male copulatory success was fecundity assessed by the father/son regression on the num- Fecundity,number of eggs laid per lifetime, was ber of females mated (Falconer, 1981). Sons were recorded for each female that a male mated. After provided with the same environment as their copulation, females remained in the carton into fathers and afforded the same opportunity to mate which they had been sequestered during copula- six females. As with fathers, none of the sons mated tion. They were provided a fresh pole bean every a seventh female within the 30 days allotted for five days. A 2 cm-wide strip of white paper towel courtship and mating although several sons did (Scott brand) was suspended from the carton lid mate with all six of the initially provided females. as a surface for oviposition. The number of yolk- For each father, one son was randomly selected bearing egg casings was recorded for each egg mass from 15 progeny (reared as described above) laid. Fecundity was determined for all females of derived from the first egg mass of the first female 26 paternal and 28 filial generation males, includ- mated. Forty father/son pairs were tested. ing 22 father/son pairs. MALE COPULATORY SUCCESS IN THE SOUTHERN GREEN STINKBUG 163 Throughout, statement of statisticalsig- 3166; n =28) males. Mean fecundity decreased nificance implies P<005. between paternal and filial generations. of copulatory success RESULTS Heritability Fecundity arid male copulatory success Significantadditive genetic variation for male cop- ulatory success is present (r=0.36; t=238; n= The fecundity of females covaried with the success 40; fig. 2). Heritability, estimated as twice the slope of their mates. In both the paternal (r =038;t= of the regression line between paternal and filial 201;n=26) and filial (r=0.52; t=256; n=28) copulatory success, was 065. generations, the lifetime fecundity of the first There was a significant positive correlation female mated by a male was significantly positively between the size of males and their success for correlated with the total number of females mated fathers (r=0.52; t=333; n=32; fig.3) but not by the male (fig. 1; The analysis is restricted to the first female mated to maintain an independence between data points that would be compromised 6.0 if the analysis incorporated multiple mates of Ca C single males). Similarly, there is a significant posi- 05.0 tive correlation between the fecundity of the last U) female mated and the number of copulations 04.0 A achieved by the male (r=051; t=290; n=27; 'a, a) 3.0 data for paternal generation only). If the average C fecundity of mates is regressed on male copulatory >a) 2.0 success, the correlation is positive for both paternal C) Ca (r=0.29; t=133;P>005; n =22)and filial gen- 1.0• erations (r =074;t= 467;P <005;n =20).
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