Adaptiveness of sex ratio control by the pupal parasitoid Itoplectis naranyae (Hymenoptera: Ichneumonidae) in response to host size

Adaptiveness of sex ratio control by the solitary parasitoid wasp Itoplectis naranyae (Hymenoptera: Ichneumonidae) in response to host size was studied, by examining whether differential effects of host size on the fitness of resulting wasps are to be found between males and females. The offspring sex ratio (male ratio) decreased with increasing host size. Larger hosts yielded larger wasps. Male larvae were less efficient in consuming larger hosts than female larvae. No significant interaction in development time was found between parasitoid sex and host size. Larger female wasps lived longer than smaller females, while longevity of male wasps did not increase with increasing wasp size. Smaller males were able to mate either with small or with large females, while larger males failed to mate with small females. Larger female wasps had a greater number of ovarioles and mature eggs at any one time than smaller females, although the number of eggs produced per host-feeding was not influenced by female wasps. Thus, the differential effect of host size on the fitness of males and females exists in I. naranyae. The basic assumption of the host-size model was therefore satisfied, demonstrating that sex ratio control by I. naranyae in response to host size is adaptive.     

Host-size-dependent sex ratio in a parasitoid wasp

Charnov's host-size model explains parasitoid host-size-dependent sex ratio as an adaptive consequence when there is a differential effect of host size on the offspring fitness of parasitoid males versus females. This article tests the predictions and the assumptions of the host-size model. The parasitoid wasp Pimpla nipponica Uchida (Hymenoptera: Ichneumonidae) laid more female eggs in larger or fresher host pupae when choice among hosts of different sizes or ages was allowed. Then, whether an asymmetrical effect of host size and age on the fitness of females versus males existed in P. nipponica was examined. Larger or fresher host pupae yielded larger wasps. Larger females lived longer, whereas male size did not influence male longevity. Large males mated successfully with relatively large females but failed with small females, whereas small males could mate successfully either with small or with large females. Thus, small-male advantages were found, and this held true even under male–male competition. Ovariole and egg numbers at any one time did not differ among females of different sizes. Larger females attained higher oviposition success and spent less time and energy for oviposition in hosts. Larger females produced more eggs from a single host meal. Taken together, females gained more, and males lost more, by being large. Host size and age thus asymmetrically affected the fitness of offspring males versus females through the relationships between host size or hast age and wasp size, which means the basic assumption of the host-size model was satisfied. Therefore, sex ratio control by P. nipponica in response to host size and age is adaptive. Received: November 13, 1998 / Accepted: January 18, 1999     

Body size and fitness relation in male and female Diaeretiella rapae

A strong relationship exists between body size and fitness in parasitoids. However, it is unclear whether the relationship is symmetric or asymmetric in males and females. The present study investigated the body size and fitness relationship in Diaeretiella rapae emerged from small and large nymphs of cabbage aphid Brevicoryne brassicae. A positive relationship existed between the size of the aphid host and growth of parasitoid larva developing in it. The fitness gain in males and females was not proportionate to their body size gain. Females mated with larger males produced 10?% more female offspring than females mated with smaller males. However, females that developed in large hosts produced 62?% more offspring (total male and female) than the females emerged from smaller hosts. The findings suggest that the number of offspring and the progeny sex ratio were affected by the body size of both male and female D. rapae.     

Non-random Mating in the Dance Fly Empis borealis: The Importance of Male Choice

In the dance-fly Empis borealis (Diptera, Empididae), females form swarms to which males, carrying a nuptial gift, come for mating. We examined whether males or females were choosy and/or competed for mates. First, measurements of the size relationships between copulating males and females, nuptial gifts and the swarming females from different swarms were assessed. Second, male visiting time in differently sized female swarms was recorded. Larger (wing-length) females participated disproportionately in copulations in each swarm, but not for the population at large. Female mating status (virgin/non-virgin) or proximity to oviposition (egg size) did not influence the likelihood of copulation. No assortative mating pattern was found: male size and size of nuptial gift did not correlate with size of the mating female. The time spent by males in swarms increased with the number of females present and it took longer when males left a swarm without copulation than when doing so. Male visiting time per female was negatively correlated with number of females in swarms. Males more often left smaller than larger swarms without mating. We conclude that E. borealis males discriminate among females but find no evidence for male competition or for female choice. It is still a question to what degree females compete for males.     

Testing predictions for the evolution of lekking in the sandfly, Lutzomyia longipalpis

We carried out a combination of field and laboratory experiments to investigate two possible mechanisms promoting the evolution of lekking in the sandfly, Lutzomyia longipalpis. Males in this species lek on or near hosts that females visit to mate and feed. To test whether the distribution of resources, lek size and female number were positively correlated (a prediction of hotspot models) we manipulated the availability of potential lek sites by varying the numbers of chickens, Gallus gallus domesticus, held in field cages. We then investigated the distribution of males and females across leks formed within these cages and measured males to assess the relationship between male size and lek size. The number of males attracted to a lek increased with the number of hosts present and on average males on larger leks had longer abdomens. The number of females was positively correlated with the number of males present, but the average number of females per male decreased with increasing lek size. In a second experiment, we introduced single females to laboratory leks of different sizes, and compared female latency to mate. Contrary to an explicit prediction of the black hole model, lek size had a direct effect on a female's latency to mate. Females at larger leks encountered and copulated with males sooner than those at smaller aggregations, but the length of time a pair spent courting did not vary with lek size. Our results suggest that the black hole model is unlikely to explain the evolution of lekking in this species. The data instead support the hotspot model as a possible mechanism promoting lekking in L. longipalpis but suggest that female preference for larger leks may need to be invoked to obtain the degree of aggregation observed.     

DIFFERENCES IN BIOMASS ALLOCATION TO REPRODUCTIVE AND VEGETATIVE STRUCTURES OF MALE AND FEMALE PLANTS OF A DIOECIOUS,PERENNIAL HERB,SILENE ALBA (MILLER) KRAUSE

The pattern of biomass allocation of males and females and the sex ratio and growth characteristics of plants from three seed-size classes in Silene alba were investigated in a greenhouse study. Seed size significantly affected adult plant size and flower production of both male and female plants, but there was no significant difference in the proportion of males and females emerging in three seed-size categories. Male and female plants differed in the proportion of total biomass allocated to vegetative and reproductive structures and these differences were consistent across all seed-size categories. Males allocated a greater proportion of their biomass to flowers than did females. Female reproductive effort was dependent upon the percentage of flowers producing mature capsules. Only females with greater than 20% fruit set have a higher reproductive expenditure than males. Consequently, female expenditure is potentially greater than males, but is spread out over a longer portion of the growing season. This difference in the timing of reproductive expenditures by males and females allows females to allocate more biomass to growth during the early flowering period and may therefore account for the common pattern in herbaceous perennial dioecious species in which adult females are larger than adult males.     

Host-feeding and acceptance by a parasitic wasp (Hymenoptera: Ichneumonidae) as influenced by egg load and experience in a patch

Optimal host selection models based on dynamic programming predict that the physiological state of a foraging insect, i.e. egg load, energy reserves etc., influences behavioral decisions. To test this prediction, the effect of physiological state on host acceptance of the ectoparasitic wasp Agrothereutes lanceolatus was investigated. Female wasps in plastic cups (regarded as patches) were presented with hosts, and their responses to the hosts were continuously observed. After observations, the wasps were dissected and the number of mature and immature eggs they carried were counted. The results showed that behavioral decisions by the female wasps were influenced by mature egg load, but not by wasp size or immature egg load. Hence the wasps with higher egg loads were more likely to oviposit. The number of hosts previously encountered in a patch (i.e. wasp experience) also had an independent effect on females' host acceptance, indicating that female informational state was updated during foraging in that patch. Female wasps host-fed only when mature egg load approached zero. Concurrent host-feeding was not observed. Parasitoid survival was almost zero when parasitoid eggs were transferred onto hosts that were fed upon, indicating that concurrent host-feeding could cause a high degree of offspring mortality. These three results supported the assumption and prediction of optimal host-feeding models. Parasitoid host selection and host-feeding are discussed in the context of recent models.     

Size-dependent mating preference of the male fiddler crab Austruca perplexa

In many animals, females prefer large males to small males, which allow large males to be choosier than small males when selecting a mate. We investigated the courtship intensity of small- and large-sized male fiddler crabs (Austruca perplexa) by examining their claw-waving rates (waves/min) towards small- and large-sized females. We found that large males showed a greater preference for large females by producing more waves/min towards them, whereas small males did not show any apparent preference for either large or small females. Moreover, the waving rate of large males was positively correlated with female size, but there was no correlation between waving rate and female size in small males. These results indicate that large males in a population become choosier and show strong mate choice, which is most likely due to their greater preference among females.     

Influence of Conspecific Presence, Experience, and Host Quality on Oviposition Behavior and Clutch Size Determination in Anastrepha ludens (Diptera: Tephritidae)

Females of some insect species adjust the number of ovipositions and clutch size adaptively depending on conspecific density and probably experience. In a series of three experiments, we examined the effect of the presence of conspecifics, host quality, and oviposition experience on oviposition behavior and clutch size determination by females of the polyphagous fruit fly Anastrepha ludens (Diptera: Tephritidae). In the first experiment, we determined that grouped (eight females per cage) A. ludens females tended to visit and oviposit in more hosts than did solitary females probably as a result of stimulation by the presence of conspecifics. We also determined that females with previous oviposition experience visited and oviposited in more hosts than inexperienced ones. Importantly, when females were grouped, we observed significantly more landings on unoccupied hosts (i.e., devoid of flies) than on occupied ones (i.e., with at least one fly on it). However, oviposition experience, and not female density, was the most important factor affecting clutch size. Naive females deposited larger egg clutches than experienced ones. In the second experiment, we found that oviposition experience and host quality (i.e., clean fruit or fruit covered with a host marking pheromone [HMP] extract), influenced clutch size and the decision of females to defend or not defend the host. Clutch size and number of fights were greater on clean than on HMP-marked hosts. In the third experiment, we observed that host quality (i.e., size) played a significant role with regard to the number of female fights, host marking behavior, and clutch size. Specifically, females fought and dragged their aculeus longer on small- and medium-sized hosts than on large ones. But this behavior varied according to whether females were kept alone or grouped. Clutch size was greatest in the largest hosts. Considering all the above, we believe that the observed increase in ovipositional activity by grouped A. ludens females can be attributed to competition through mutual interference and not social facilitation as has been reported in other tephritid species.     

Interference of asexual and sexual reproduction in the green hydra

The green hydra, Hydra viridissima, has three sexes: hermaphrodite, male, and female. I investigated the reproductive strategies of the green hydra and the relationship between asexual budding and sexual reproduction. The proportion of mature individuals in the asexually reproducing population increased with increasing temperature. Sexual reproduction did not interrupt asexual budding in hermaphrodites or males; sexual reproduction did interrupt asexual budding in females. Sexual reproduction also resulted in exponential population growth during the reproductive season. The number of asexual buds on each parental individual was positively correlated with the parental individual size in asexual individuals and in males. The same positive correlation was found between the number of testicles and the size of males. These correlations reflect a common tendency in asexual and sexual reproduction: larger parental individuals have a greater number of propagules or gametes. No correlation was found between size and buds or size and gonads in hermaphrodites; hermaphrodites had at most one asexual bud and were significantly larger than males, females, and asexual individuals. The larger size of hermaphrodites supports the hypothesis that producing both female and male gonads is more energetically costly than producing only one type of gamete (gonochorism).     

Effects of female and male size on female mating and remating decisions in a bean beetle

The body sizes of individuals of the choosing and chosen sexes in a mate choice may affect sequential mating of females. We examined the effects of the body sizes of females and their mates on attributes of female first mating, and the effects of body sizes of females and their previous and potential future mates on female remating in the adzuki bean beetle, Callosobruchus chinensis. Large- and small-sized adults were derived from larvae reared under conditions of low and high density in a bean, respectively. The speed of first mating of large females was not affected by the size of courting males, whereas small females initiated mating more rapidly when courted by small males. The remating probability of large females was not affected by first male size, whereas small females that mated first with smaller males were more likely to remate. These data suggest that pre- and post-copulatory female choices for male size depend on the female’s size, and the small females might be more willing to copulate with smaller males but prefer larger males to sire their offspring after copulation. A possible explanation for this preference is that small females may suffer greater harm from copulating with larger males.     

The determinants of sexual segregation in the scalloped hammerhead shark,Sphyrna lewini

Synopsis Female scalloped hammerhead sharks move offshore at a smaller size than do males to form schools composed primarily of intermediate size female sharks. This movement results in smaller females feeding more on pelagic prey than do males and with greater predatory success. It is contended that this change in habitat causes females to grow more rapidly to reproductive size. Intermediate size females grow at a more rapid rate than males. Female scalloped hammerhead sharks mature at a size larger than males. For many elasmobranch species, females: (1) occupy a different habitat, (2) grow more rapidly prior to maturity and continue growth following maturation, (3) feed on different prey with increased feeding success, and (4) reproduce at a size larger than males. It is suggested that female segregation increases fitness, resulting in more rapid growth for the former sex. The females reach maturity at the larger size necessary to support embryonic young, yet similar age to males, matching the female reproductive lifetime to that of males.     

蜡皮蜥的两性异形和繁殖输出

为研究蜡皮蜥(Leiolepis reevesii)两性异形和繁殖输出,于2002、2003年4月下旬从海南乐东一种群捕获423头蜡皮蜥。经检测得到繁殖雌体的最小体长为89．0mm,据此判定≥89．0mm的个体为性成熟。研究结果表明：①蜡皮蜥具有两性异形,雄性大于雌性且具有较大的头部。成体雄性头长和头宽随体长的增长速率大于雌性,幼体头长和头宽随体长的增长速率无显著的两性差异。以性别和年龄(成、幼体)为因子的双因子ANOVA比较两性头长和头宽与体长的回归剩余值发现,雄性头部大于雌性,幼体头部相对大于成体。②饲养于实验室的母体中有42头于2002、2003年5月22日～7月16日产出正常卵,这些繁殖雌体具有年产多窝卵的潜力。窝卵数和卵重的变异系数分别为0．18和0．13,前者变异度大于后者。窝卵数、窝卵重和卵重均与母体体长无关。卵重与相对生育力之间无显著的负相关性,表明蜡皮蜥缺乏卵数量与卵大小之间的权衡。相对窝卵重与母体体长呈显著的负相关,表明较小的母体具有相对较大的繁殖输出。因雌体繁殖会滞缓其生长,小母体具有相对较大的繁殖输出,至少部分地解释了雌性蜡皮蜥的成体为什么个体较小。     

Asymmetric size effect of sexes on reproductive fitness in an aphid parasitoid Aphidius ervi (Hymenoptera: Aphidiidae)

Most studies on size–fitness relationships focus on females and neglect males. Here, we investigated how body size of both sexes of an aphid parasitoid, Aphidius ervi Haliday, affected the reproductive fitness. Reproductive fitness was generally positively correlated with body size for both sexes in this species. Large individuals of both sexes had greater longevity, large males fathered more progeny, and large females had higher fecundity, parasitism, and greater ability in host searching and handling. We demonstrated in this study that size effects of males and females were asymmetric on different reproductive fitness parameters. With increasing body size females gained more than males in longevity and fecundity while males gained more than females in the number of female progeny. Regardless of female size, large males sustained a female-biased population longer than small males. These results suggest that male body size should also be considered in the quality control of mass-rearing programs and the evaluation of parasitoid population growth.     

Effect of the primary host for production of both sexes on the mating interaction in an autoparasitoid species

Encarsia sophia (Girault and Dodd) is an autoparasitoid in the hymenopteran family Aphelinidae. The females develop as primary parasitoids on whitefly nymphs (primary hosts), whereas the males develop as hyperparasitoids on their own species or on other primary parasitoid species (secondary hosts). The autoparasitoids not only parasitise whiteflies but also kill them with strong host-feeding capacity. In this study, female and male E. sophia were reared on the primary hosts Trialeurodes vaporariorum and Bemisia tabaci ‘Q’, and the host-feeding and parasitism of wasps on both whitefly species were determined for the four possible different mating combinations: (i) E. sophia females reared on B. tabaci (ESF-BT) mated with E. sophia males from B. tabaci (ESM-BT), (ii) E. sophia females reared on T. vaporariorum (ESF-TV) mated with E. sophia males from T. vaporariorum (ESM-TV), (iii) ESF-BT mated with ESM-TV, and (iv) ESF-TV mated with ESM-BT. ESF-TV mated with ESM-TV killed the largest percentage of whitefly nymphs through host feeding. The ESF-TV with larger body size mating with larger ESM-TV killed more whitefly nymphs through host feeding than those mating with smaller ESM-BT. Whether B. tabaci or T. vaporariorum were used as hosts, ESF-TV mated with ESM-TV and ESM-BT and ESF-BT mated with ESM-BT significantly parasitised more whitefly nymphs than ESF-BT mated with ESM-TV. In general, ESF-BT mated with ESM-TV killed significantly fewer whitefly nymphs through parasitism and host feeding than the other three mating combinations on both whitefly species. These results indicated that the performance of autoparasitoids on insect pests was not only dependent on females but was also affected by mating with males from different primary host species.     

Mutual mate choice by mountain pine beetles: size‐dependence but not size‐assortative mating

1. Mutual mate choice may be rare, occurring when both sexes invest heavily in reproduction, mating opportunities are abundant, and individuals differ in quality. 2. Mountain pine beetles, Dendroctonus ponderosae (Curculionidae: Scolytinae) appear to meet the conditions for mutual mate choice. We introduced males to females in breeding sites and observed the occurrence and speed of a male entering a female's gallery. We tested for consequences of mutual mate choice, namely condition‐dependent choosiness and assortative mating. 3. Males were more likely to enter a female's gallery when the gallery was in a smaller tree with less resin production and when the gallery was larger. Female body size and condition did not influence the probability of entry. Larger males were less likely to enter a gallery than were smaller males, probably because of size‐dependent choosiness rather than physical limitations. 4. Small males took longer to enter galleries of large females than of small females, whereas large males entered as quickly into galleries of large females as small females. This suggests size‐dependent choosiness by females. 5. No assortative mating by body size was detected, probably because males appeared to choose on the basis of female‐associated resources rather than on female traits.     

THE EFFECTS OF BODY SIZE ON MATING FEATURES IN A FIELD POPULATION OF THE HERMAPHRODITIC SEA HARE APLYSIA KURODAI BABA, 1937 (GASTROPODA: OPISTHOBRANCHIA)

Mating behaviour of the hermaphroditic sea hare Aplysia kurodaiBaba was observed in the field to investigate the effects ofbody size on various mating features. Compared with small individuals,large gastropods mated with more partners both as males andas females, and also with larger partners (size-assor-tativemating). Large gastropods also mated more frequently as femalesbut not as males, and tended to be selected as female matingpartners more often than small ones by neighbouring individualsbut not as male partners. All these results suggest some matingadvantage of large individuals over small ones as females, probablybecause of the preference by male partners for large femalepartners. At the same time, the results suggest that the matingadvantage of large size as males may be limited, if any. Thepreference for large female partners is also consistent withthe fact that their mating was longer than that of small ones.The greater advantage of large size in mating as females, thanas males, means that sexual selection at mate determinationacts more strongly on female size than on male size and thatthe direction of total sexual selection at mating is probablyreversed in this hermaphrodite (Received 27 November 1995; accepted 8 February 1996)     

Floral display, pollinator visitation and reproductive success in the dioecious perennial herb Wurmbea dioica (Liliaceae)

Floral traits that increase attractiveness to pollinators are predicted to evolve through selection on male function rather than on female function. To determine the importance of male-biased selection in dioecious Wurmbea dioica, we examined sexual dimorphism in flower size and number and the effects of these traits on pollinator visitation and reproductive success of male and female plants. Males produced more and larger flowers than did females. Bees and butterflies responded to this dimorphism and visited males more frequently than females, although flies did not differentiate between the sexes. Within sexes, insect pollinators made more visits to and visited more flowers on plants with many flowers. However, visits per flower did not vary with flower number, indicating that visitation was proportional to the number of flowers per plant. When flower number was experimentally held constant, visitation increased with flower size under sunny but not overcast conditions. Flower size but not number affected pollen removal per flower in males and deposition in females. In males, pollen removal increased with flower size 3 days after flowers opened, but not after 6 days when 98% of pollen was removed. Males with larger flowers therefore, may have higher fitness not because pollen removal is more complete, but because pollen is removed more rapidly providing opportunities to pre-empt ovules. In females, pollen deposition increased with flower size 3 days but not 6 days after flowers opened. At both times, deposition exceeded ovule production by four-fold or more, and for 2 years seed production was not limited by pollen. Flower size had no effect on seed production per plant and was negatively related to percent seed set, implying a tradeoff between allocation to attraction and reproductive success. This indicates that larger flower size in females is unlikely to increase fitness. In both sexes, gamete production was positively correlated with flower size. In males, greater pollen production would increase the advantage of large flowers, but in females more ovules may represent a resource cost. Selection to increase flower size and number in W. dioica has probably occurred through male rather than female function. Received: 15 June 1997 / Accepted: 12 February 1998     

Ejaculate investment and attractiveness in the stalk‐eyed fly,Diasemopsis meigenii

The phenotype‐linked fertility hypothesis proposes that male fertility is advertised via phenotypic signals, explaining female preference for highly sexually ornamented males. An alternative view is that highly attractive males constrain their ejaculate allocation per mating so as to participate in a greater number of matings. Males are also expected to bias their ejaculate allocation to the most fecund females. We test these hypotheses in the African stalk‐eyed fly, Diasemopsis meigenii. We ask how male ejaculate allocation strategy is influenced by male eyespan and female size. Despite large eyespan males having larger internal reproductive organs, we found no association between male eyespan and spermatophore size or sperm number, lending no support to the phenotype‐linked fertility hypothesis. However, males mated for longer and transferred more sperm to large females. As female size was positively correlated with fecundity, this suggests that males gain a selective advantage by investing more in large females. Given these findings, we consider how female mate preference for large male eyespan can be adaptive despite the lack of obvious direct benefits.     

Breeding Strategies in Females of the Parasitoid Wasp Spalangia endius: Effects of Mating Status and Size

Does the mating status or body size of a female parasitoid wasp affect her host size choice or propensity to burrow? In Spalangia endius, using smaller hosts appears to reduce a female's cost of parasitization but not her son's fitness. However, virgin females, which produce only sons, did not preferentially parasitize smaller hosts. Mated females also showed no host size preference. Mated females burrowed more than virgins in the presence of hosts, although not in their absence. Burrowing may reduce a mated female's harassment from males, and not burrowing may increase a virgin female's chance of mating because males avoid burrowing. Mating did not increase female longevity. Greater female size increased the offspring production of mated females burrowing for hosts but not in the absence of burrowing and not in virgin females. A female's size had no significant effect on whether her first drill attempt was on a large or a small host or on the duration of her successful drills.