Food intake of fruit-feeding butterflies: evidence for adaptive variation in proboscis morphology

Adult butterflies feed from a variety of substrates and have appropriate adaptations. We examined proboscis morphology in a community of fruit-feeding butterflies (Nymphalidae) in a tropical forest in Uganda. These data were supplemented with behavioural observations and measurements of intake rate on natural and artificial substrates. We found no sexual dimorphism in proboscis morphology even though puddling behaviour is usually performed by males. Two main feeding techniques could be distinguished on the basis of behaviour and morphology: the piercing technique, typically found in Charaxinae, and the sweeping technique employed by both Nymphalinae and Satyrinae. These techniques, distinguished in previous studies, are described in more detail and their relative efficiencies are discussed in the context of sexual dimorphism, food-choice and life history evolution.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 86 , 333–343.     

Life history strategies and habitat templets of tropical butterflies in north-eastern Australia

Three multivoltine species of satyrine butterflies in the genus Mycalesis (Lepidoptera: Nymphalidae) are narrowly sympatric in the wet–dry tropics of north-eastern Australia. They show a range of ecological strategies and adaptations associated with contrasting habitats and varying selective pressures. Two abiotic factors, namely favorability (the reciprocal of seasonal adversity) and predictability (broadly the reciprocal of disturbance), were examined as potential environmental selective forces in shaping their life histories. Comparison of several key life history traits of the wet-season form revealed that the life histories of each species corresponded well with their habitat characteristics. M. perseus, which lives in habitats which are less favorable (i.e. adverse) and more unpredictable (i.e. temporary), shows many traits of an r-type strategy: smaller size, faster development, earlier maturation, higher fecundity, smaller egg size, and rapid population increase. By contrast, M. sirius and M. terminus, which live in more favorable and predictable (i.e. permanent) habitats, have many life history attributes and other characteristics in common which link them closer to K-type strategies. The only discrepancy is lower potential reproductive effort of M. perseus, which may be accounted for in terms of an evolutionary trade-off, such as with dispersal or dormancy. Other correlates associated with the M. perseus life history tactic include higher sex-size dimorphism, greater dispersal ability, better tolerance to adverse conditions, stronger phenotypic variation, greater degree of polyandry, and a more flexible breeding strategy. The life history patterns of these species are discussed in the context of evolutionary life history models, particularly the Southwood–Greenslade habitat templet.     

Preferences and Food Quality of Fruit‐Feeding Butterflies in Kibale Forest,Uganda1

Many butterflies in tropical forests feed on fruits that have fallen to the forest floor. This substrate differs in many ways from floral nectar, and therefore fruit‐feeding butterflies are expected to possess adaptations for efficient foraging, choice, and ingestion of their food. Differences in food quality and in spatial and temporal availability are also likely to have led to life history evolution. Here we describe the sugar and nitrogen content of fruits that butterflies feed on in a tropical forest in Uganda, and measure the attractiveness of these fruits to the local butterfly fauna together with the role that decay plays in the attraction. These data are supplemented with feeding observations at fruit falls in the forest. Our results show that (1) fruits contain significant and variable concentrations of sugar and nitrogen, and constitute a nutritious food source for butterflies in tropical forests; (2) fruit‐feeding butterflies use cues from the fruits and fermentation products to locate their food; (3) different classes of fruit‐feeding butterflies may vary in their preferences for certain fruits, and differ in their ability to find preferred food; and (4) fruit choice is not strongly correlated with attractiveness or nutrient content. The results are discussed in the light of the evolution of food searching and life history strategies.     

Genetic relationships between Oeneis urda and O. mongolica (Nymphalidae: Lepidoptera)

The species status of Oeneis urda (Eversmann) and O. mongolica (Oberthür) has been argued based on morphological characters. Reexamination of their major morphological characters has shown a slight differentiation in the two species. Sequences of three mitochondrial genes (COI, ND6, and ND1) and one nuclear region (internal transcribed spacer 2, ITS2) from two O. urda populations (Yangyang and Mt. Hanla) and one O. mongolica population (Uljin) were performed for phylogenetic and population genetic inferences. Sharing of identical sequences in the ND6 gene and ITS2, minimal sequence divergence in the COI and ND1 genes, and phylogenetically undividable sequence types in all mitochondrial genes and ITS2 suggest genetic continuity between the two species. Nevertheless, significant F_ST estimates (P < 0.05) were found for the COI gene in comparisons between Yangyang (O. urda) and Uljin (O. mongolica), between Yangyang (O. urda) and Mt. Hanla (O. urda), and between Uljin (O. mongolica) and Mt. Hanla (O. urda) populations. These F_ST estimates, along with other gene‐based analyses collectively suggest isolation of the two species at some point in the past, but incomplete separation between the two species on the mainland (Yangyang and Uljin) and biogeographically forced isolation of the O. urda population on Mt. Hanla collectively appear to complicate species status of these two species that were once further clearly separated.     

The evolution and genetics of sexually dimorphic ‘dual’ mimicry in the butterfly Elymnias hypermnestra

Sexual dimorphism is a major component of morphological variation across the tree of life, but the mechanisms underlying phenotypic differences between sexes of a single species are poorly understood. We examined the population genomics and biogeography of the common palmfly Elymnias hypermnestra, a dual mimic in which female wing colour patterns are either dark brown (melanic) or bright orange, mimicking toxic Euploea and Danaus species, respectively. As males always have a melanic wing colour pattern, this makes E. hypermnestra a fascinating model organism in which populations vary in sexual dimorphism. Population structure analysis revealed that there were three genetically distinct E. hypermnestra populations, which we further validated by creating a phylogenomic species tree and inferring historical barriers to gene flow. This species tree demonstrated that multiple lineages with orange females do not form a monophyletic group, and the same is true of clades with melanic females. We identified two single nucleotide polymorphisms (SNPs) near the colour patterning gene WntA that were significantly associated with the female colour pattern polymorphism, suggesting that this gene affects sexual dimorphism. Given WntA''s role in colour patterning across Nymphalidae, E. hypermnestra females demonstrate the repeatability of the evolution of sexual dimorphism.     

Dynamics of mate-searching behaviour in a hilltopping butterfly, Lasiommata megera (L.): the effects of weather and male density

By studying a hilltopping population of the satyrine Lasiommata megera in southern Sweden, the effects of density and meteorological conditions on mate-searching behaviour were investigated. Lasiommata megera males switched between stationary and vagrant behaviour, and the behaviour adopted was correlated to meteorological conditions; more males were stationary at low temperatures, low irradiation levels and high wind speeds than vice versa. Body temperature measurements indicated that these factors were important in deciding the body temperature and thereby the flight activity of males. Male density did not significantly influence mate-locating behaviour. The tendency of males to hilltop and their distribution around the hill were strongly influenced by weather. The hilltopping behaviour of this species did not adhere to an all-or-none pattern, but instead males, as well as the stations they used, were found at increasingly higher elevations on the hill with increasing temperatures and decreasing wind speeds. Moreover, males tended to use the lee side and sun-exposed side of the hill. Females, of which the majority were already mated, also showed this distribution around the hill, but preferred lower elevations than males. Release experiments did not reveal any propensity of mated females to fly uphill. Such a propensity was shown by released virgin females.