Diversity in structural ultraviolet coloration among female sulphur butterflies (Coliadinae, Pieridae)

In some species of sulphur butterflies (Pieridae: Coliadinae) females as well as males display bright structural reflectance on their dorsal wing surfaces, although comparatively little attention has been paid to this coloration in females. We examined the spectral properties of female dorsal coloration and scale structure in three species of sulphurs for which published images show bright UV reflectance in females: the Neotropical Anteos clorinde and two species of Indo-Australian Eurema, E. hecabe and E. candida. In A. clorinde and E. hecabe, female UV reflectance is iridescent and produced by thin film interference in a system of ridges and lamellae, as it is in conspecific males. Female A. clorinde exhibit the same spatial distribution and chromaticity of UV reflectance as seen in males, but the UV reflectance in female E. hecabe is much smaller in area compared to that of conspecific males and is both less bright and less chromatic than observed in males. In contrast, UV reflectance in E. candida females is diffuse, and arises from a lack of pterin pigments in the wings, which permits a broad-band scattered reflection to be seen. This is the mechanism that is known to produce bright UV reflectance in females of the confamilial whites. Our results highlight the diversity of UV reflectances and underlying mechanisms in sulphurs and suggest multiple evolutionary pathways leading to this diversity in female sulphur butterflies.     

Sexual differences in tailwind drift compensation in Phoebis sennae butterflies (Lepidoptera: Pieridae) migrating over seas

One prediction derived from optimal migration theory is thatmigrating animals that maximize their flight distance on agiven amount of energy will decrease their airspeed in a tailwindand increase it in a headwind. To test this in a migratingbutterfly, I followed male and female cloudless sulfur butterfliesPhoebis sennae (Pieridae) migrating from Colombia toward Panamaover the Caribbean Sea. P. sennae headed westerly over the Caribbean Sea in the morning and then turned southeasterly tohead downwind in the afternoon. Changes in heading and trackdirections of P. sennae were not related to changes in theposition of the solar azimuth. As predicted from optimal migrationtheory, flight velocities of females decreased in a tailwindto minimize energy consumption. However, males did not showany compensation for tailwinds. Females are minimizing energyconsumption, whereas males may be minimizing the time to reachthe destination site in order to maximize matings with newlyarrived or newly emerged females. Orientation of females changedbefore that of males, presumably because their greater reproductiveload imposed greater flight costs and limited flight fuels.     

New Delias butterflies from the Solomon Islands (Lepidoptera: Pieridae)

Abstract Four new taxa of Delias from the Solomon Islands are described. Two are races of a new species, D-messalina sp.n., which may be closely related to D.nigrina Fabricius and is distributed over three islands. The nominotypical subspecies, D.m.messalina subsp.n., occurs in Bougainville while D.m.orientalis subsp.n. is found on Santa Ysabel and Guadalcanal. The other two, D.alberti guava subsp.n. from Bougainville and D.alberti tetamba subsp.n. from Santa Ysabel, represent new subspecies of D.alberti Rothschld. Couplets for the inclusion of the new taxa in the standard key in Talbot are provided. In addition, a list of Solomon Islands Delias is given, together with brief distributional data.     

DNA barcodes identify Central Asian Colias butterflies (Lepidoptera,Pieridae)

A majority of the known Colias species (Lepidoptera: Pieridae, Coliadinae) occur in the mountainous regions of Central-Asia, vast areas that are hard to access, rendering the knowledge of many species limited due to the lack of extensive sampling. Two gene regions, the mitochondrial COI ‘barcode’ region and the nuclear ribosomal protein RpS2 gene region were used for exploring the utility of these DNA markers for species identification. A comprehensive sampling of COI barcodes for Central Asian Colias butterflies showed that the barcodes facilitated identification of most of the included species. Phylogenetic reconstruction based on parsimony and Neighbour-Joining recovered most species as monophyletic entities. For the RpS2 gene region species-specific sequences were registered for some of the included Colias spp. Nevertheless, this gene region was not deemed useful as additional molecular ‘barcode’. A parsimony analysis of the combined COI and RpS2 data did not support the current subgeneric classification based on morphological characteristics.     

The evolutionary history of the ‘alba’ polymorphism in the butterfly subfamily Coliadinae (Lepidoptera: Pieridae)

Polymorphisms are common in the natural world and have played an important role in our understanding of how selection maintains multiple phenotypes within extant populations. Studying the evolutionary history of polymorphisms has revealed important features of this widespread form of phenotypic diversity, including its role in speciation, niche breadth, and range size. In the present study, we examined the evolutionary history of a ubiquitous colour polymorphism in the sulphur butterflies (subfamily: Coliadinae) termed the ‘alba’ polymorphism. We investigated the origin and stability of the ‘alba’ polymorphism using ancestral state reconstruction analysis. Our results indicate that the ancestor of the Coliadinae was polymorphic and that this polymorphism has undergone repeated transitions to monomorphism. Repeated loss of polymorphism suggests that the ‘alba’ polymorphism may be relatively unstable over evolutionary time. These results provide a framework for future studies on the origin and maintenance of the ‘alba’ polymorphism and guide the direction of future hypotheses. We discuss these results in light of current understandings of how the ‘alba’ polymorphism is maintained in extant populations.     

DNA barcoding and species delimitation of butterflies (Lepidoptera) from Nigeria

Molecular Biology Reports - Accurate identification of species is a prerequisite for successful biodiversity management and further genetic studies. Species identification techniques often require...     

A mitochondrial-DNA-based phylogeny for some evolutionary-genetic model species of Colias butterflies (Lepidoptera, Pieridae)

We study the phylogenetic relationships among some North American Colias ("sulfur") butterflies, using mitochondrial gene sequences (ribosomal RNA, cytochrome oxidase I+II) totaling about 20% of the mitochondrial genome. We find that (1) the lowland species complex shows a branching order different from earlier views; (2) several montane and northern taxa may be more distinct than in earlier views; (3) one morphologically conservative Holarctic assemblage, C. hecla, is differentiated at the molecular-genetic level into at least three taxa which occupy distinct positions in the phylogeny and are sisters to diverse other taxa. These conclusions, constituting phylogenetic hypotheses, are supported by parsimony, maximum-likelihood, and Bayesian reconstruction algorithms. They are tested formally, by interior branch tests and paired-site tests, against alternative hypotheses derived from conventional species and subspecies naming combinations. In all cases our hypotheses are supported by these tests and the conventional alternatives are rejected. The "barcoding" subset of cytochrome oxidase I sequence identifies only some of the taxa supported by our full data set. Comparison of genetic divergence values among Colias taxa with those among related Pierid butterflies suggests that species radiations within Colias are comparatively younger. This emerging Colias phylogeny facilitates comparisons of genetic polymorphism and other adaptive mechanisms among taxa, thereby connecting micro- and macro-evolutionary processes.     

Thermoregulatory significance of wing melanization in Pieris butterflies (Lepidoptera: Pieridae): physics,posture, and pattern

Summary Pieris butterflies use a novel behavioral posture for thermoregulation called reflectance basking, in which the wings are used as solar reflectors to reflect radiation to the body. As a means of exploring the thermoregulatory significance of wing melanization patterns, I examine the relation of basking posture and wing color pattern to body temperature. A mathematical model of the reflectance process predicts certain combinations of dorsal wing melanization pattern and basking posture that maximize body temperature. Laboratory experiments and field observations show that this model correctly predicts qualitative differences in the relation of body temperature to basking posture based on differences in the extent of dorsal melanization on the wing margins, both between species and between sexes within species of Pieris. This is the first demonstration in insects that coloration of the entire wing surface can affect thermoregulation. Model and experimental results suggest that, in certain wing regions, increased melanization can reduce body temperature in Pieris; this effect of melanization is exactly the opposite of that found in other Pierid butterflies that use their wings as solar absorbers. These results are discussed in terms of the evolution of wing melanization pattern and thermoregulatory behavior in butterflies.     

Müllerian mimetic radiation of Delias butterflies (Lepidoptera: Pieridae) in Bali and Timor

Mimicry rings are present among Delias butterflies, and those butterflies are also considered to be mimetic models of other lepidopteran insects; however, experimental evidence for their unpalatability to predators is limited. In Bali and Timor, a total of three mimicry rings of Delias species are present; particularly, male and female D. lemoulti join different rings in Timor. The present study examined the unpalatability of Delias in Bali and Timor to the caged avian predator Pycnonotus aurigaster. The birds ate eight Delias species in similar numbers, and ate the palatable butterfly Mycalesis horsfieldii much more frequently than Delias butterflies. The result suggests that the three mimicry rings of Delias species in Bali and Timor are Müllerian rather than Batesian. Based on previous findings on their phylogenetic relationships, the Müllerian mimicry rings of Delias in Bali and Timor are suggested to have emerged through the convergent evolution and phylogenetic constraints of wing color patterns. In the D. hyparete species group, mimetic radiation may have occurred between Bali and Timor.     

Revised systematics and higher classification of pierid butterflies (Lepidoptera: Pieridae) based on molecular data

The butterfly family Pieridae comprises approximately 1000 described species placed in 85 genera, but the higher classification has not yet been settled. We used molecular data from eight gene regions (one mitochondrial and seven nuclear protein‐coding genes) comprising a total of ~6700 bp from 96 taxa to infer a well‐supported phylogenetic hypothesis for the family. Based on this hypothesis, we revise the higher classification for all pierid genera. We resurrect the tribe Teracolini stat. rev. in the subfamily Pierinae to include the genera Teracolus, Pinacopteryx, Gideona, Ixias, Eronia, Colotis and most likely Calopieris. We transfer Hebomoia to the tribe Anthocharidini and assign the previously unplaced genera Belenois and Dixeia to the subtribe Aporiina. Three lineages near the base of Pierinae (Leptosia, Elodina and Nepheronia + Pareronia) remain unplaced. For each of these, we describe and delineate new tribes: Elodinini Braby tribus nova, Leptosiaini Braby tribus nova and Nepheroniini Braby tribus nova. The proposed higher classification is based on well‐supported monophyletic groups and is likely to remain stable even with the addition of more data.     

Evolution of larval food plant associations in Delias Hübner butterflies (Lepidoptera: Pieridae)

A review of larval food plants of the genus Delias is presented. Larvae specialize primarily on aerial‐stem and root hemiparasites (“mistletoes”) in the order Santalales. Although butterfly food plant associations have been recorded for only a small proportion of the genus (28 species or ～11%, representing 12/24 species‐groups), available data suggest that the family Loranthaceae is used most frequently (77%), followed by the Santalaceae sensu stricto (14%) and Viscaceae (8%). With the possible exception of Euphorbiaceae (1%), almost all non‐mistletoe records are considered to be erroneous and, in most cases, probably represent the mistletoe host tree on which the larvae sometimes pupate. Of the eight major clades recognized in Delias, food plants have been recorded for six of these, although the majority of records (89%) are for three clades (hyparete, belladonna, nigrina). Optimization of the larval food plant data in the context of recent phylogenetic hypotheses for both butterflies and plants revealed little evidence of cospeciation at the higher systematic levels. The most parsimonious reconstruction was an origin of larval feeding on Loranthaceae, followed by at least six independent colonizations to Santalaceae + Viscaceae. In contrast to related pierids in the Aporiina associated with mistletoes in which further shifts from aerial‐stem mistletoes to distantly related plants (e.g. host trees parasitized by mistletoes) have facilitated differentiation at the generic level, there is no firm evidence to indicate that such secondary, monomorphic shifts have evolved in Delias. However, larvae of D. henningia (pasithoe group of belladonna clade) from Palawan and Luzon, the Philippines, appear to be polymorphic, feeding on both Loranthaceae and Euphorbiaceae.     

Thermal ecology of Pieris butterflies (Lepidoptera: Pieridae): a new mechanism of behavioral thermoregulation

Summary I document a new mechanism for behavioral thermoregulation, not previously described in animals, called reflectance basking. This behavior, described here for Pieris butterflies, involves the use of the wings as solar reflectors that reflect solar radiation onto the body to increase body temperature. Results show that Pieris require thoracic (body) temperature. between 29° and 40° C in order to take off and fly, and achieve these elevated temperatures by basking. Diurnal patterns of population flight activity are closely correlated with patterns of body temperature during basking. Behavioral studies indicate that 1) Pieris orient to solar radiation, 2) they use thermoregulatory postures consistent with reflectance basking, and 3) they do not use the basking postures found in other Pierid butterflies (i.e., the Coliadinae). There are consistent differences in wing angles used in reflectance basking between Pieris in different subgenera. Results are discussed with respect to thermoregulation and wing color in other Pierid butterflies, and suggest that a re-evaluation of the functional significance of melanization in Pieris is needed.     

Hypersensitivity reaction of Brassica nigra L. (Cruciferae) kills eggs of Pieris butterflies (Lepidoptera: Pieridae)

Summary Some individual plants of the mustard Brassica nigra in lowland California kill eggs of the Crucifer-specialist herbivores Pieris rapae and P. napi by producing a necrotic zone at the base of the egg, thereby apparently desicating it. This is a typical hypersensitivity reaction, but to an atypical stimulus. The eggs can be rescued by maintaining them in a saturated atmosphere. Attempts to demonstrate a bacterial or fungal agent associated with the reaction were unsuccessful.     

Phylogenetic relationships among genera of danaine butterflies (Lepidoptera: Nymphalidae) as implied by morphology and DNA sequences

Cladistic relationships among genera and subtribes of Danaini (the milkweed butterflies) were inferred by analysis of data combined from five sources: morphology of adults and immature stages, and DNA sequences from three gene regions. The results corroborate and greatly increase support for prior hypotheses based on morphology alone. A new index summarizing incongruence among data partitions, the Partition Congruence Proportion (PCP), is introduced. The significance of the inferred pattern of phylogenetic relationships for comparative chemical ecology of milkweed butterflies is briefly discussed.     

Phylogenetic systematics of Colotis and associated genera (Lepidoptera: Pieridae): evolutionary and taxonomic implications

We investigated the genetic diversity and phylogenetic placement of the butterflies in the genus Colotis and eight related pierid genera using sequence information from two mitochondrial and two nuclear genes. To establish the status of species, we initially barcoded 632 specimens representative of all genera and most species and subspecies in those genera. A subset was then selected for phylogenetic analysis where additional gene regions were sequenced: 16S rRNA (523 bp), EF‐1α (1126 bp) and wg (404 bp). DNA barcode results were largely congruent with the traditional classification of species in the Colotis group, but deep splits or lack of genetic divergence in some cases supported either species‐level differentiation or synonymy. Despite using information from four genes, the deeper nodes in our phylogeny were not strongly supported, and monophyly of the ‘Colotis group’ and the genera Colotis and Eronia could not be established. To preserve the monophyly of Colotis, we revive the genus Teracolus for three outlying species previously in Colotis (i.e. Colotis eris, Colotis subfasciatus and Colotis agoye), as well as the genus Afrodryas for Eronia leda. The position of Calopieris is unresolved although it appears to be well outside the molecular variation in Colotis (s.l.). A dispersal/vicariance analysis suggested that major diversification in Colotis (s.str.) occurred in Africa with subsequent dispersal to India and Madagascar.     

Phylogenetic relationships and divergence times among New World monkeys (Platyrrhini, Primates)

Orthologous sequences of six nuclear genes were obtained for all recognized genera of New World monkeys (Primates: Platyrrhini) and outgroups to evaluate the phylogenetic relationships and to estimate divergence times. Phylogenetic relationships were reconstructed by maximum parsimony, maximum likelihood, and Bayesian approaches. All methods resolved with 100% branch support genus-level relationships, except for the grouping of Aotus as a sister taxa of Cebus and Saimiri, which was supported by low bootstrap percentages and posterior probability. All approaches depict three monophyletic New World monkey families: Atelidae, Cebidae, and Pitheciidae; also within each family, all approaches depict the same branching topology. However, the approaches differ in depicting the relationships of the three families to one another. Maximum parsimony depicts the Atelidae and Cebidae as sister families next joined by the Pitheciidae. Conversely, likelihood and Bayesian phylogenetic trees group families Atelidae and Pitheciidae together to the exclusion of Cebidae. Divergence time estimations using both local molecular clock and Bayesian approaches suggest the families diverged from one another over a short period of geological time in the late Oligocene-early Miocene.     

Performance and host preference of Ascia monuste (Lepidoptera,Pieridae)

Experiments were conducted to determine the performance and host preference of Ascia monuste using kale (Brassica oleraceae, var. Acephala) and mustard (B. juncea). These plants differ significantly in water and nitrogen content, with mustard having larger amounts of water and kale larger amounts of nitrogen. The performance results confirmed that kale is a better food source than mustard for the species, even when eggs were collected on mustard leaves in the field. However, when eggs were collected on mustard, the kale nutritive value was lower than the nutritive value obtained when eggs were collected on kale leaves. Furthermore, the results of oviposition preference obtained in the field and in the laboratory have shown a preference for kale, indicating the presence of a positive correlation with performance. In contrast to the data about oviposition preference, there was no immature feeding preference. These results indicate that host selection occurs during the oviposition process. Furthermore, it is possible that the high abundance of kale cultivated in the region studied and the nutritional quality of this plant are two factors that influence the positive relationship between oviposition preference and performance for A. monuste.     

Survival of refrigerated Tatochila butterflies (Lepidoptera: Pieridae) as an indicator of male nutrient investment in reproduction

Summary Tatochila butterflies show prolonged survival in cold (2° C) storage. Mating significantly reduces survival in males, suggesting that nutrients may be directed into spermatophores or maintenance under storage conditions; this provides a potential index of male nutrient investment in reproduction.