A Survey of Eyespot Sexual Dimorphism across Nymphalid Butterflies (original) (raw)

Female Bicyclus anynana butterflies choose males on the basis of their dorsal UV-reflective eyespot pupils

… of the Royal …, 2005

Sexual and natural selection pressures are thought to shape the characteristic wing patterns of butterfly species. Here we test whether sexual selection by female choice plays a role in the maintenance of the male wing pattern in the butterfly Bicyclus anynana. We perform one of the most extensive series of wing pattern manipulations in butterflies, dissecting every component of the 'bulls-eye' eyespot patterns in both ventral and dorsal wing surfaces of males to test the trait's appeal to females. We conclude that females select males on the basis of the size and brightness of the dorsal eyespot's ultraviolet reflecting pupils. Pupil absence is strongly selected against, as are artificially enlarged pupils. Small to intermediate (normal sized) pupils seem to function equally well. This work contradicts earlier experiments that suggest that the size of dorsal eyespots plays a role in female choice and explains why male dorsal eyespots are very variable in size and often have indistinct rings of coloration, as the only feature under selection by females seems to be the central white pupil. We propose that sexual selection by female choice, rather than predator avoidance, may have been an important selective factor in the early stages of eyespot evolution in ancestral Lepidopteran lineages.

MaleBicyclus anynanaButterflies Choose Females on the Basis of Their Ventral UV-Reflective Eyespot Centers

Journal of insect science, 2019

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.

Quantifying wing pattern sexual dimorphism in the butterfly Anartia amathea roeselia Eschscholtz (Lepidoptera: Nymphalidae)

International Journal of Fauna and Biological Studies, 2015

The present study searched for morphological markers in the wings of Anartia amathea roeselia butterfly. This species can be used as a model species due to its natural abundance in SE Brazil and Neotropics. It is an excellent tool for analysis of phenotypical traits and in studies of population genetics. In this research, wing pattern of both sexes were described and quantified in a sample of 812 butterflies considering sexual dimorphism related to size and colour. The results revealed that approximately 55% of the marker spots on the dorsal surface of wings presented little (< 2.5%) or no variation. Therefore, only ten of 22 spots were used as markers in the analysis, with some spots being sexually dimorphic. The female size was bigger than males as well as other Nymphalidae species. The results showed that the colour wing pattern of A. a. roeselia had enough differences to separate individuals of both sexes and morphological markers that can be used to study the role of natural...

Polyphenism of visual and chemical secondary sexually-selected wing traits in the butterfly Bicyclus anynana: How different is the intermediate phenotype?

PLoS ONE, 2019

Polyphenism is a type of phenotypic plasticity supposedly adaptive to drastic and recurrent changes in the environment such as seasonal alternation in temperate and tropical regions. The butterfly Bicyclus anynana shows polyphenism with well-described wet and dry seasonal forms in sub-Saharan Africa, displaying striking morphological, physiological and behavioural differences in response to higher or lower developmental temperatures. During the seasonal transition in the wild, the intermediate phenotype co-occurs with wet and dry phenotypes. In this study, we aimed to characterize the secondary sexually-selected wing traits of the intermediate form to infer its potential fitness compared to wet and dry phenotypes. Among the previously described wing morphological traits, we first showed that the area of the fifth eyespot on the ventral hindwing is the most discriminant trait to identify wet, dry and intermediate phenotypes in both sexes. Second, we characterized the intermediate for...

Sexual Size Dimorphism in the Color Pattern Elements of Two Mimetic Heliconius Butterflies

Sexual dichromatism and sexual dimorphism of body size are reasonably well studied in butterflies. Sexual size dimorphism of color pattern elements, however, is much less explored. The object of this study is Heliconius, a genus of butterflies well known for the coevolution between mate color preferences and mimicry. Given the sexual role of wing coloration, we investigated the existence of sexual size dimorphism in the wing color elements of a mimetic pair—Heliconius erato phyllis Fabricius and Heliconius besckei Ménétriés—and analyzed the allometric patterns of these traits. Correlation between size of elements in the dorsal and ventral wing surfaces were also estimated. In both species, three out of four elements were larger in males, but the nondimorphic element was not the same. With regard to the allometric patterns, our most important finding was that smaller males of one species have proportionally larger yellow bars. This is the first study specifically concerning quantitative sexual dimorphism in the coloration of this well-known genus of butterflies and it opens new prospects to investigate sex-related natural selection and sexual selection of color pattern elements.

Male disguised females: costs and benefits of female-limited dimorphism in a butterfly

1. In polymorphic species, two or more discrete phenotypes co-occur simultaneously. Sex-limited polymorphism is a particular case of polymorphism, in which several discrete morphs coexist within one of the two sexes only. Several hypotheses were proposed to explain the existence and the maintenance of sex-limited polymorphism in insects: (i) the morphs have similar fitness, such as similar survival and expected fecundity, and their frequencies vary randomly (i.e. the null hypothesis); (ii) harassment by males is reduced towards the less common female morph, in this case andromorph females (i.e. the male mimicry and learned mate recognition hypotheses); (iii) morphs differ in predation risk (i.e. the predation hypothesis); or (iv) morphs differ in thermoregulation ability (i.e. the thermoregulation hypothesis). 2. Field observations and experiments were employed to compare the relative support of these hypotheses using dimorphic females of the bog fritillary butterfly. Differences were detected between morphs in survival, fecundity, harassment by males, predation pressure and thermal properties, thereby rejecting the null hypothesis. 3. The lifestyle of both morphs is associated with different costs and benefits, with advantages in daily survival and precocious emergence for the gynomorph females, and advantages in fecundity, predation and male harassment for the andromorph females. Besides, as the bog fritillary butterfly is protandrous (i.e. males emerge before females), the longer development of andromorph females puts them at risk of emerging when all the males are dead. The results raise the question as to which mechanisms control the ontogenetic pathways driving the production of the two morphs (i.e. genetic polymorphism or phenotypic plasticity).

Wing Morphology and Body Design in Opsiphanes and Caligo Butterflies Match the Demands of Male Mating Displays (Lepidoptera: Nymphalidae)

Annals of the Entomological Society of America, 2020

There is noticeable variation in male mate-seeking behavior among species of Brassolini butterflies. Males of Opsiphanes Doubleday and Caligo Hübner (Lepidoptera, Nymphalidae) species perform crepuscular displays along forest edges and in light gaps. While male Opsiphanes perform aerial displays, Caligo males perch and wait for receptive females. A comparison of five species of each genus suggests that male display behavior is associated with, and has likely influenced the evolution of their wing attributes and body design. Opsiphanes males have higher wing aspect ratio and more distal centroid position than congeneric females, suggesting that the energetic demands of aerial displays led to sexual dimorphism in wing morphology. In contrast, male and female Caligo generally showed similar wing morphology, which possibly results from the lower energy expenditure of perching behavior when compared with active flight. Likely due to a genetic correlation between sexes, female Opsiphanes ...

The relationship between eyespot shape and wing shape in the butterfly Bicyclus anynana: A genetic and morphometrical approach

Journal of Evolutionary Biology, 1997

The African butterfly, Bicyclus anynana, normally possesses circular eyespots on its wings. Artificial selection lines, which express ellipsoidal eyespots on the dorsal surface of the forewing, were used to investigate correlated changes in wing shape. Morphometric analysis of linear wing measurements and wing scale counts provided evidence that eyespot shape was correlated with localised shape changes in the corresponding wing-cell, with overall shape changes in the wing, and with the density/arrangement of scales around the eyespot area.