Nectar sugar and amino acid preferences of Battus philenor (Lepidoptera, Papilionidae)

Abstract.

• 1 Butterflies of Battus philenor were tested for their preferences for nectar sugars and amino acids in an outdoor cage experiment.
• 2 The butterflies clearly preferred both sucrose and fructose over glucose. They also preferred sucrose over fructose.
• 3 No other preferences were found to be statistically significant, although male butterflies tended to prefer a plain sugar solution over a sugar solution containing a mixture of amino acids: females consumed both of these solutions in almost equal proportions.
• 4 The results are discussed with respect to nectar composition of butterfly pollinated flowers, flower preferences, physiological and reproductive aspects of butterflies.

     

Contemporary patterns in a historical context: phylogeographic history of the pipevine swallowtail,Battus philenor (Papilionidae)

Abstract We examined mitochondrial DNA (mtDNA) variation in pipevine swallowtail butterflies ( Battus philenor ) from throughout its extant range to provide a historical, phylogeographical context for ecological studies of the disjunct population in California. We evaluate current hypotheses regarding host plant use, behavior, and mimetic relationships of B. philenor populations and generate alternative hypotheses. Compared to populations throughout the rest of the species' range, California populations are ecologically distinct in that they lack mimics, lay significantly larger clutches of eggs, and exclusively use a unique, endemic larval host plant. Analysis of molecular variance, tests of population differentiation, and nested clade analysis of mtDNA variation indicate that, despite low levels of population genetic structure across the species' range, there is evidence of recent range expansion from presumed Pleistocene refuge(s) in southeastern North America. Colonization of California appears to have been a recent event. This phylogeographic investigation also suggests that the evolution of life-history adaptations to a novel larval host has occurred rapidly in California and the lack of mimics in California may be attributable to the recency of colonization.     

Are gardens effective in butterfly conservation? A case study with the pipevine swallowtail, <Emphasis Type="Italic">Battus philenor</Emphasis>

Gardens with nectar sources and larval host plants have been proposed to stem the decline in butterfly abundance caused by habitat loss. However, no study has provided evidence that gardens benefit butterflies. We examined the use of natural sites and gardens in the San Francisco bay area by the butterfly, Battus philenor. We found that natural sites were more likely to attract adult B. philenor, received more oviposition, and had higher juvenile survival than gardens sites. Butterflies were more likely to be present in gardens with established populations of the host plant, Aristolochia californica, growing in the sun. Battus philenor are unlikely to visit gardens with host plants planted within the past 7 years. Gardens between the ages of 8–40 years received oviposition, but did not always support completion of larval development of B. philenor. In gardens with host plants over 40 years of age, B. philenor consistently survived from egg to the adult stage. Natural enemy induced mortality of eggs did not differ between garden and natural sites, but overall egg survival was lower in gardens than at natural sites. It is unlikely that gardens serve as 'refugia' for B. philenor in years when populations in natural sites experience low survival or low fecundity. Even in gardens capable of supporting larvae to maturity, the density of eggs and survival rates were lower than in natural populations of the host plant suggesting that gardens were not optimal habitats. Therefore, without evidence that juvenile abundance and survival rates in gardens matches or exceeds that in natural sites, it is most likely that gardens act as population sinks for B. philenor.     

Chemical defence of Battus philenor larvae against attack by the parasitoid Trogus pennator

Abstract 1. Parasitoids in the genus Trogus (Hymenoptera: Ichneumonidae) attack the larvae of swallowtail butterflies (Lepidoptera: Papilionidae). Only two of the three major tribes of the subfamily Papilioninae are attacked although species of all three tribes commonly occur together. The tribe Troidini is relatively free of parasitoids of any kind, and it has been proposed that the aristolochic acids sequestered by troidines protect them from parasitism.
2. The responses of T. pennator (Fabricius) to the sympatric troidine Battus philenor (Linnaeus) were examined. Three hypotheses that could explain why this wasp does not parasitise B. philenor were considered. (1) Battus philenor does not produce compounds used by the wasp to locate hosts. (2) The larval integument contains compounds that deter attack. (3) The parasitoid offspring cannot survive in B. philenor .
3. The first hypothesis was not supported as the frass of B. philenor larvae was found to act as a searching arrestant comparable to the frass of a host species.
4. The second hypothesis was supported. The B. philenor larvae were rejected when the wasps examined them using their antennae, and ethanolic washes of B. philenor cuticle deterred attack by T. pennator when applied to otherwise acceptable host larvae. Bioassays of fractions of the ethanolic wash and of pure aristolochic acids established that aristolochic acids were at least partly responsible for the deterrent effect.
5. The third hypothesis was also supported. Larvae of B. philenor attacked by the parasitoids developed into butterflies.
6. These results indicate that both behavioural and physiological barriers, the former attributable at least in part to sequestered compounds and the latter of unknown mechanism, prevent T. pennator from parasitising B. philenor .     

No geographic variation in thermoregulatory colour plasticity and limited variation in heat‐avoidance behaviour in Battus philenor caterpillars

Phenotypic plasticity can help organisms cope with variation in their current environment, including temperature variation, but not all environments are equally variable. In the least variable or extreme environments, plasticity may no longer be used. In this case, the plasticity could be lost altogether, or it could persist with either the same or an altered reaction norm, depending on factors such as the plasticity's costs. In the pipevine swallowtail caterpillar (Battus philenor), I tested for changes in two forms of heat‐avoidance plasticity, colour change and refuge‐seeking behaviour, across the species’ range in the United states, including the cooler eastern parts of its range where colour change has not been observed and is unlikely to be needed. I found that both heat‐avoidance behaviour and colour change persisted in all surveyed populations. Indeed, the reaction norm for colour change remained nearly unaltered, whereas the threshold for refuge‐seeking only changed slightly across populations. These results suggest that the costs of these plastic traits are low enough for them to be maintained by whatever minimal gene flow the population receives. I show that plasticity can be maintained unaltered in populations where it is not used and discuss the potential consequences of this persistence for both the ecology and evolution of plasticity.     

Pollen amino acids—an additional diet for a nectar feeding butterfly?

The increase of the amino acid concentration over different time intervals in artificial nectar (i.e., a sucrose solution) due to pollen contamination was investigated in four Californian plant species (Aesculus californica, Amsinckia lunaris, Brodiaea pulchella, Carduus pycnocephalus), which are important nectar resources for a Californian colony of the butterflyBattus philenor as well as for other insects. The increase of the amino acid concentration in the medium is different in all four species and seems to be determined by a variety of factors including permeability of the pollen grain wall and presence or absence of pores. The results suggest a passive diffusion process of the free pollen amino acids into the medium rather than an active release. Implications from the experiments forBattus philenor and for other nectar feeding pollinators are discussed. A possible complementary effect of free pollen and nectar amino acids is proposed for plant species in which pollen is likely to be knocked into nectar by their flower visitors. A possible evolutionary pathway from nectar feeding butterflies such asBattus philenor to the complex derived pollen feeding habit in theHeliconius butterflies is proposed.     

Warning Signal Efficacy: Assessing the Effects of Color,Iridescence, and Time of Day in the Field

Warning coloration deters predators from attacking distasteful or toxic prey. Signal features that influence warning color effectiveness are not well understood, and in particular, we know very little about how effective short‐wavelength and iridescent colors are as warning color elements in nature and how warning signal effectiveness changes throughout the day. We tested the effect of these factors on predation risk in nature using specimens of the distasteful pipevine swallowtail butterfly, Battus philenor. B. philenor adults display both iridescent blue and diffusely reflecting orange components in their warning signal. We painted B. philenor wings to create five different model types: all‐black, only‐iridescent‐blue, only‐orange, iridescent‐blue‐and‐orange (intact signal), and matte‐blue‐and‐orange. We placed 25 models in each of 14 replicate field sites for 72 h and checked for attacks three times each day. Model type affected the likelihood of attack; only‐orange models were, the only model attacked significantly less than the all‐black model. Iridescence did not enhance or decrease warning signal effectiveness in our experiment because matte‐blue‐and‐orange models were attacked at the same rate as iridescent‐blue‐and‐orange models. Time of day did not differentially affect model type. Video recordings of attacks revealed that insectivorous birds were responsible. The results of this experiment, when taken with previous work, indicate that the response to blue warning coloration is likely dependent on predator experience and context, but that iridescence per se does not affect warning signals in a natural context.     

Comparative processing of allelochemicals in the Papilionidae (Lepidoptera)

Within the family Papilionidae (Lepidoptera), species display a broad range of feeding patterns, from oligophagy on a single hostplant family to polyphagy on over a dozen families. Accompanying this diversity of feeding strategies is a diversity of physiological mechanisms for processing hostplant allelochemicals. Studies on members of this family as well as other Lepidoptera suggest that oligophagy is associated with high activity, in addition to high substrate specificity, of detoxicative enzymes.     

Larger clutches of chemically defended butterflies reduce egg mortality: evidence from Battus philenor

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