Males Become Choosier in Response to Manipulations of Female Wing Ornaments in Dry Season Bicyclus anynana Butterflies.

Male investment towards reproduction is substantial in some species, and this leads to the evolution of choosy males. Male choice is often directed towards female phenotypes that are good indicators of fecundity such as body size, age, or virgin status, and often acts in the same direction as fecundity selection. In insects, only a few examples exist where male choice is directed towards female ornaments as proxies Butler of female quality. Here, we use dry season males of the sex-role reversed butterfly species Bicyclus anynana to test for differences in male choosiness and investment towards females of varying attractiveness using ornament-manipulations. Male reproductive investment in this species is in the form of a nuptial gift, a spermatophore, given to females upon mating. Males were placed in cages with either wild-type ornamented females or with nonornamented females (center of the dorsal forewing eyespots painted over), and time to mating, duration of mating, and longevity of males and females after a single mating were measured. Ornament manipulations consisted of blocking the UV-reflective scales in the center of the dorsal forewing eyespots of females, a known sexual ornament. Males displayed lower latency to mate and longer mating durations with ornamented females. The longer mating duration did not, however, translate in the transfer of a nuptial gift that increased female longevity or reduced male longevity. Instead, we propose that longer mating durations with ornamented females may represent increased mate guarding behavior or increased sperm transfer.

Ng SY ，Bhardwaj S ，Monteiro A 《JOURNAL OF INSECT SCIENCE》

Male Bicyclus anynana Butterflies Choose Females on the Basis of Their Ventral UV-Reflective Eyespot Centers.

Butterflies often use their dorsal and ventral wing color patterns for distinct signaling functions. Color patterns on hidden dorsal wing surfaces are often used in sexual signaling, while exposed ventral patterns are often used to ward off predator attacks. At rest, however, part of the ventral forewings are often hidden by the hindwings, allowing individuals to also use the patterns on this wing surface for sexual signaling. Here, we test this hypothesis in Bicyclus anynana (Butler, Lepidoptera, Nymphalidae) butterflies by first determining the degree of sexual dimorphism in ventral forewing patterns, focusing on the eyespots, from both wet and dry season forms, and then testing the role of the larger ventral forewing eyespots of dry season females in male mate choice. We also test male investment in reproduction. We show that ventral forewing UV-reflective eyespot centers, in addition to dorsal forewing eyespot centers previously examined in this species, play a role in sexual signaling as males preferentially mated with females with their ventral eyespot centers intact instead of blocked with black paint. This male preference, however, did not translate into a detectable higher reproductive investment via a single mating toward ornamented females. This study provides an example of how ventral forewing patterns, often hidden by hindwings, are used in sexual communication, in this case by females to attract males.

Huq M ，Bhardwaj S ，Monteiro A 《JOURNAL OF INSECT SCIENCE》

Developmental plasticity in sexual roles of butterfly species drives mutual sexual ornamentation.

Current explanations for why sexual ornaments are found in both sexes include genetic correlation, same sex competition, and mutual mate choice. In this study, we report developmental plasticity in mating behavior as induced by temperature during development in the butterfly Bicyclus anynana. Males and females reciprocally change their sexual roles depending on their larval rearing temperatures. This switch is correlated with a change in mating benefits to females and costs to males. The discrete seasonal environments, wet season and dry season, are known to produce the two developmental forms and as a consequence impose alternating, symmetrical patterns of sexual selection, one season on male ornaments, the following season on female ornaments. Thus, reciprocal selection through time may result in mutual sexual ornamentation.

Prudic KL ，Jeon C ，Cao H ，Monteiro A ... -  《-》

Biased learning affects mate choice in a butterfly.

Early acquisition of mate preferences or mate-preference learning is associated with signal diversity and speciation in a wide variety of animal species. However, the diversity of mechanisms of mate-preference learning across taxa remains poorly understood. Using the butterfly Bicyclus anynana we uncover a mechanism that can lead to directional sexual selection via mate-preference learning: a bias in learning enhanced ornamentation, which is independent of preexisting mating biases. Naïve females mated preferentially with wild-type males over males with enhanced wing ornamentation, but females briefly exposed to enhanced males mated significantly more often with enhanced males. In contrast, females exposed to males with reduced wing ornamentation did not learn to prefer drab males. Thus, we observe both a learned change of a preexisting mating bias, and a bias in ability to learn enhanced male ornaments over reduced ornaments. Our findings demonstrate that females are able to change their preferences in response to a single social event, and suggest a role for biased learning in the evolution of visual sexual ornamentation.

Westerman EL ，Hodgins-Davis A ，Dinwiddie A ，Monteiro A ... -  《-》

Rearing Temperature Influences Adult Response to Changes in Mating Status.

Rearing environment can have an impact on adult behavior, but it is less clear how rearing environment influences adult behavior plasticity. Here we explore the effect of rearing temperature on adult mating behavior plasticity in the butterfly Bicyclus anynana, a species that has evolved two seasonal forms in response to seasonal changes in temperature. These seasonal forms differ in both morphology and behavior. Females are the choosy sex in cohorts reared at warm temperatures (WS butterflies), and males are the choosy sex in cohorts reared at cooler temperatures (DS butterflies). Rearing temperature also influences mating benefits and costs. In DS butterflies, mated females live longer than virgin females, and mated males live shorter than virgin males. No such benefits or costs to mating are present in WS butterflies. Given that choosiness and mating costs are rearing temperature dependent in B. anynana, we hypothesized that temperature may also impact male and female incentives to remate in the event that benefits and costs of second matings are similar to those of first matings. We first examined whether lifespan was affected by number of matings. We found that two matings did not significantly increase lifespan for either WS or DS butterflies relative to single matings. However, both sexes of WS but not DS butterflies experienced decreased longevity when mated to a non-virgin relative to a virgin. We next observed pairs of WS and DS butterflies and documented changes in mating behavior in response to changes in the mating status of their partner. WS but not DS butterflies changed their mating behavior in response to the mating status of their partner. These results suggest that rearing temperature influences adult mating behavior plasticity in B. anynana. This developmentally controlled behavioral plasticity may be adaptive, as lifespan depends on the partner's mating status in one seasonal form, but not in the other.

Westerman E ，Monteiro A 《PLoS One》