R. I. Vane-Wright

The Biology of butterflies

Background and Identification Butterflies are in the order Lepidoptera which are insects with scaled wings. Moths are also in the order Lepidoptera. Butterflies are invertebrates, which means that they do not have a backbone, instead they have an exoskeleton, a shell that encases their soft body and protects their vital organs. Butterflies undergo metamorphosis, so that the immature and adult forms are very different.

It's time to work together and stop duplicating conservation efforts ...

We strongly support initiatives to produce clear, efficient and practical goals for conservation to guide biodiversity planners and decision-makers in governments, agencies, conventions and non-governmental organizations (NGOs). However, as things stand there is only limited consensus on global conservation priorities at international level. We believe that the time is now right for scientists and practitioners to work together to develop a commonly adopted blueprint for action.

The systematic position of Antirrhea and Caerois, with comments on the classification of the Nymphalidae (Lepidoptera)

Based on a Wagner tree analysis of ninety‐two characters (eighty‐five larval, one egg, six adult), the nymphalid butterfly genera Antirrhea and Caerois are demonstrated to be the closest relatives of the genus Morpho. Accordingly, Antirrhea and Caerois are formally transferred from the Satyrinae to the Morphinae. Without these two genera, the Morphinae (sensu Ehrlich) is at best a paraphyletic group. During the study, fourteen nymphalid genera were treated as potential outgroups. The analysis suggests that many currently accepted higher taxa within the Nymphalidae are probably untenable: the Satyrinae and Nymphalinae (sensu Ehrlich) are both probably polyphyletic; the Biinae (sensu Miller) must be abandoned, being polyphyletic; the Charaxidae (sensu Rydon), although probably monophyletic, appear to form a group subordinate to part of the ‘Satyrinae’; and Apatura does not cluster with the ‘Nymphalinae’, but appears to form the sister‐group of the ‘Satyrinae’ (less Antirrhea and Caerois) plus the Charaxinae. Re‐analyses of reduced data sets, in which potentially homoplasious larval head‐horn and adult wing venational characters were eliminated, leaves these conclusions essentially unaltered. The authors suggest that a solution to the seemingly intractable problem posed by nymphalid higher classification can be sought by the application of cladistic analysis to a large data set gathered from all developmental stages, with special emphasis on detailed comparative larval morphology.

Independent gene phylogenies and morphology demonstrate a malagasy origin for a wide-ranging group of swallowtail butterflies.

Abstract Madagascar is home to numerous endemic species and lineages, but the processes that have contributed to its endangered diversity are still poorly understood. Evidence is accumulating to demonstrate the importance of Tertiary dispersal across varying distances of oceanic barriers, supplementing vicariance relationships dating back to the Cretaceous, but these hypotheses remain tentative in the absence of well‐supported phylogenies. In the Papilio demoleus group of swallowtail butterflies, three of the five recognized species are restricted to Madagascar, whereas the remaining two species range across the Afrotropical zone and southern Asia plus Australia. We reconstructed phylogenetic relationships for all species in the P. demoleus group, as well as 11 outgroup Papilio species, using 60 morphological characters and about 4 kb of nucleotide sequences from two mitochondrial (cytochrome oxidase I and II) and two nuclear (wg and EF‐1α) genes. Of the three endemic Malagasy species, the two that are formally listed as endangered or at risk represented the most basal divergences in the group, while the more common third endemic was clearly related to African P. demodocus. The fifth species, P. demoleus, showed little differentiation across southern Asia, but showed divergence from its subspecies sthenelus in Australia. Dispersal‐vicariance analysis using cladograms derived from morphology and three independent genes indicated a Malagasy diversification of lime swallowtails in the middle Miocene. Thus, diversification processes on the island of Madagascar may have contributed to the origin of common butterflies that now occur throughout much of the Old World tropical and subtemperate regions. An alternative hypothesis, that Madagascar is a refuge for ancient lineages resulting from successive colonizations from Africa, is less parsimonious and does not explain the relatively low continental diversity of the group.

THE CLADISTICS OF AMAURIS BUTTERFLIES: CONGRUENCE, CONSENSUS AND TOTAL EVIDENCE

Abstract— Two data sets for 10 species of African milkweed butterflies (Nymphalidae, Danainae: one Danaus, two Tirumala, seven Amauris) have been analysed cladistically, separately and in combination. One data set comprised 32 morphological characters, the other comprised 68 chemical compounds from male scent organs. Analysed separately, the two data sets produced six similar but non‐identical minimum‐length solutions. Analysed together, the combined data set of 100 characters produced a single minimum‐length tree, identical to one of the three solutions for the morphological data set. The combined data produced a more informative result than congruence comparisons based on strict or combinable component consensus analysis. These results, together with re‐analysis of a morphological data set for all 15 species of Amauris (which produced 12 minimum‐length solutions), permit increased resolution of the existing classification of this Afrotropical genus, including the formal recognition of two subgenera, Amauris (Amauris) Hübner, and Amauris (Amaura) Geyer (stat. rev.). The fit of uniquely derived, unreversed chemical characters to the tree raises the possibility that stepwise additive evolution of semiochemicals may have occurred during cladogenesis of these mimetic butterflies. The implications for chemoecology and speciation are briefly discussed.

Indifferent Philosophy versus Almighty Authority: on consistency, consensus and unitary taxonomy

Abstract The ability of the taxonomic community to heed Charles Godfrays wake‐up call to create ‘unitary’ taxonomic systems and make them available on the internet is hampered by real difficulties over achieving taxonomic consistency, and a cultural reluctance amongst systematists to embrace consensus. This paper explores these issues by examining the taxonomic history of an African milkweed butterfly, Amauris damocles sensu lato. Recent differences of opinion over the classification of this insect relate to fundamental differences in the theory and practice of systematics, from creationism through authoritarianism to cladistics, differences of a kind that will forever bedevil the distributed taxonomic system. If practical unitary schemes are to emerge, then the taxonomic community will need to adopt new ways of managing and recording taxonomic change, and develop a more responsible attitude towards the needs of others who are dependent of the primary products of systematics ‐names, and the classification schemes they symbolize.

Phylogenetic relationships of three African swallowtail butterflies, Papilio dardanus, P. phorcas and P. constantinus : a cladistic analysis (Lepidoptera : Papilionidae)

Abstract. The well‐known polymorphic mimetic African papilionid butterfly, Papilio dardanus Brown, is demonstrated by cladistic analysis to form a monophyletic group (the phorcas‐group) with two other African swallowtails, P.phorcas Cramer and P.constantinus Ward. Within this group, dardanus and phorcas are shown to be sister‐species, with constantinus as the stem. The supposed relationship of this group of three Rutaceae‐feeding species to the five members of the Canellaceae/Lauraceae‐feeding Papilio hesperus‐group (including P.nobilis Rogenhofer) is questioned. The significance of this analysis is discussed with special reference to Trimens original hypothesis on the evolution of mimetic patterns in female Papilio dardanus, the unique male patterns of phorcas and dardanus, and natural and laboratory‐produced hybrids. A pathway modelling technique is developed to represent the possible course of evolution of the phorcas‐group, in terms of speciation events and the accumulation and decay of the pattern polymorphisms.

Taxonomy, Methods of

The goal of taxonomy is to create a general reference system of biological classification. Using characters derived from sensory data gathered from individual organisms, taxonomists perform five main functions: discrimination (primary recognition of groups, or taxa), systematization (assessing the relationships among taxa), classification (summary schemes expressing the major patterns revealed), symbolization (giving unambiguous names to taxa), and identification (matching unidentified organisms to the system). Following a historical account, the article focuses on the emergence of three major methods of systematization (evolutionary systematics, phenetics, and cladistics) and its consequences on the taxonomic system.

Colour and size variation in Junonia villida (Lepidoptera, Nymphalidae): subspecies or phenotypic plasticity?

Colour pattern and size variation in the widespread Indo-Pacific Meadow Argus butterfly, Junonia villida (Fabricius, 1787), are reviewed based largely on examination of c. 1500 museum specimens from across its entire geographical range. Including villida, 14 available species-group names for this taxon are documented. The butterfly is recorded from some 200 islands or island groups. Larvae are reported to feed on 24 genera of flowering plants from 12 families. While the suggestion of H.T.G. Watkins that geographical variation in villida can be summarized by four segregates (eastern Pacific, Maluku–Papuan, Sunda–Australian, Chagos Islands) is found to have some validity, there are numerous exceptions. Based on a discussion of biological factors that may be responsible for the pattern of variation observed, it is concluded that, with the exception of J. villida chagoensis, a geographical isolate with a recognisable and constant phenotype not seen in any other population, any attempt at further formal division of this butterfly into subspecies is at present difficult or impossible to support. Reaching this conclusion has only been possible through the use of major museum collections. This leads to the perspective that variable species exhibiting seasonal polyphenism and migration, such as J. villida, need to be studied using a variety of techniques to explore their phenotypic plasticity before attempts are made to recognise numerous subspecies.

Anomalous areas and awkward ages: concerns about over-reliance on model-based biogeographical and temporal inferences

A recent molecular phylogenetic study of the cosmopolitan nymphalid butterfly genus Vanessa (Wahlberg & Rubinoff, 2011) exemplifies some potential pitfalls of algorithm-based inferences drawn from temporal and biogeographical patterns of taxonomic divergence. There has lately been a trend towards inclusion of model-based molecular clock and ancestral area estimates in phylogenetic analyses of lepidopteran taxa (e.g. Müller & Beheregaray, 2010; Price et al., 2011), and we are concerned about the potential for naive acceptance of results from such computer-based approaches. We offer the following comments in the hope of encouraging our esteemed colleagues to consider their data as well as their models.