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In theory, the selective attacks by experienced predators are expected to drive aposematic prey species towards colour monomorphism because mutant individuals with divergent colour forms will not be recognized as distasteful and thus will be targeted by predators. The tropical butterfly Danaus chrysippus represents a notable exception to this rule since it is aposematic but shows extensive colour pattern polymorphism in the region of East and Central Africa. In this paper, we present a critical review of the various hypotheses that were suggested to explain the origin and maintenance of colour polymorphism in D. chrysippus. Three different approaches to this problem were revised and criticized: first, that polymorphism is a non-adaptive consequence of past geographic isolation that resulted in the formation of a hybrid zone, second, that polymorphism represents an evolutionary escape route when an aposematic colour form is heavily loaded by batesian mimics and third, that polymorphism is maintained because D. chrysippus is subjected to spatially and temporally variable Mullerian selection as a result of recurrent population extinctions in the mimetic Acraea species, driven by the spread of the male-killing Wolbachia bacterium. It has been argued that the three hypotheses are not mutually exclusive; alternatively, each hypothesis targets a distinct problem and exploits a distinct set of key observations. The study concluded that the final solution to the long standing problem of aposematic polymorphism in D. chrysippus might be achieved through the adoption of a pluralistic approach that integrates all the three hypotheses.

The butterflies are beautiful creature of nature with great economic importance as pollinator as well as bio-indicator of environments. The present survey was conducted to determine the biology and distribution of butterfly fauna of Hazara University, Garden Campus, Mansehra, Pakistan during March-June 2012. The study area was divided into 3 quadrates, i.e., residential area, administration area and main campus. A total of 170 specimens were collected, 10 species were identified belonging to 3 different families and falling in 8 genera. The species were identified. The painted lady, Cynthia cardui (Linnaeus); blue pansy, Junonia orithya Linnaeus; and plain tiger, Danaus chrysippus (Linnaeus) were belonging to family Nymphalidae. The lime butterfly, Papilio demoleus Linnaeus and com-mon mormon, P. polytes Linnaeus were belonging to family Papilionidae. The dark clouded yellow, Colias croceus (Geoffroy); common grass yellow, Eumera hecab (Linnaeus); Murree green-veined white, Pieris ajaka Moore; green-veined white, P. napi (Linnaeus) and Bath white, Pontia daplidice (Linnaeus) were belonging to family Pieridae. The body sizes of E. hecabe and J. orithya were minimum, i.e., 1.4 cm (n = 4 - 13), however, D. chrysippus was maximum, i.e., 2.5 cm (n = 1). The wingspan of E. hecabe was minimum, i.e., 3.7 cm (n = 4), however, P. polytes was maximum, i.e., 7.5 cm (n = 1). Proper preventive measures should be taken into consideration in order to minimize the natural habitat loss, as butterfly fauna is dependent upon proper environmental conditions.

Foraging responses of seven butterfly species were determined for flower models that differed in color, size, and depth of artificial corolla. We measured frequency of visits to flowers of various colors (red, orange, yellow, purple, and white) with or without the application of honey water to flower models. We found that the relative strength of butterfly response to visual versus olfactory cues differed among butterfly species. We suggest that the seven butterfly species tested exhibit four different patterns in how visual and olfactory cues were combined as determinants of foraging importance: (1) vision was given priority over olfaction (in Cethosia cyane and Danaus chrysippus); (2) olfaction was given priority over vision (Cethosia biblis, Idea leuconoe, and Tirumala limniace); (3) olfaction and vision were equally important (Catopsilia pomona); and (4) only olfaction was used (e.g., Danaus genutia, which was not sensitive to the five colors tested but very sensitive to the smell of honey in our experiments). In tests with flower models of different sizes, the visitation frequency of C. cyane, I. leuconoe, and D. chrysippus increased with increasing corolla diameter, provided flowers were sprayed with honey water. C. cyane and D. chrysippus showed this trend because of their strong dependence on visual cues. In contrast, I. leuconoe with its large body size preferred larger flowers, it could be that butterflies need more nectar to support its large body size. Catopsilia pomona tended to visit large and middle-sized flowers because it used both vision and odor to detect food, large or middle-sized flowers could have more vision and odor stimulation than small flowers for foraging butterfly. The other three butterfly species did not show that trend because of lack of dependence on visual cues. There was no correlation between visitation frequency and corolla depth within particular butterfly species. In conclusion, the foraging strategies of butterflies are not only mainly determined by the weights given to visual and olfactory cues, but also affected by body size.

Within the region of East and Central Africa, the aposematic butterfly D. chrysippus is characterized by unusual ecology, due to the co-occurrence of polymorphism, mimicry and sex ratio distortion. In this paper, we describe a survey conducted for D. chrysippus at Khartoum, based upon morphological and molecular investigations. The goal of the study was to determine whether Sudan is a part of the presumed hybrid zone of D. chrysippus in Africa, where the butterfly shows the unusual ecology. D. chrysippus was found to display considerable colour pattern polymorphism. Only one mimetic form, the batesian mimic, female Hypolimnas misippus was recorded. The population sex ratio of D. chrysippus did not show significant deviation from the 1:1 ratio. Molecular tests for the male-killing bacteria Spiroplasma yielded negative results. The study concluded that central Sudan belongs to the hybrid zone but lacks its characteristic ecology.

Within certain regions in East Africa, the butterfly Danaus chrysippus (L.) shows female-biased population sex ratio, because of the production by some females of all-female broods, as a result of infection by maternally inherited, male-killing bacterium of the genus Spiroplasma. In this study, we describe a 3-year field survey for the population dynamics of the male-killing Spiroplasma in D. chrysippus in four independent localities, namely Uganda, Ghana, Sudan and Madagascar. The prevalence of the bacterium was found to show extensive variations at multiple scales among different sites, in various countries, seasons and years. A novel, selection-based hypothesis was suggested to explain the high variability of male-killer prevalence over space and time, based on the existence of an adaptive link between larval food-plant density and the magnitude of resource reallocation fitness advantage for the male-killer.

A 900 km2 coastal area in southern Spain was searched for both Danaus plexippus and D. chrysippus and their foodplants. A set of 63 single or mixed patches of Asclepias curassavica and Gomphocarpus fruticosus was detected and visited regularly from June 2008 to December 2009. Danaus plexippus was on the wing throughout our study period, but it was never detected in 21 patches while in 4 patches it was observed on more than 80% of our visits. Danaus chrysippus was on the wing only during summer and autumn of 2009 and it was never detected in 49 patches. The proportion of plant patches occupied by butterflies varies seasonally, increasing during summer and autumn. These results are discussed in relation to the origin, persistence and conservation of both species in Europe.

On two occasions, on opposite sides of the African continent (Cape Coast, Ghana, and Dar es Salaam, Tanzania), high adult population densities in the polymorphic butterfly Hypolimnas misippus (a presumed mimic of Danaus chrysippus) were followed by linkage disequilibrium in combinations of fore- and hindwing colour patterns. On both occasions, disequilibrium was caused by significant changes in morph frequencies favouring rarer and more mimetic forms. Recaptures were too few for analysis at Dar, although the changes there took place within a single generation and must have been the result of differential survival. Recapture rate data and survival rate estimates at Cape Coast support the hypothesis that selective predation was responsible, as does the observation of synchronous linkage disequilibrium at Dar in the model D. chrysippus, indicating parasitic mimicry. There was clear selection for the perfection of mimicry for forewings at Dar and for hindwings at Cape Coast. Disequilibrium is also reported for two other sites, Legon (Ghana) and Boksburg (South Africa) and, in all four sites, it was associated with an increase in the most mimetic forms. New chemical evidence is presented to support the contention that D. chrysippus is a defended model. Although all the evidence leads to the conclusion that H. misippus is a Batesian mimic of D. chrysippus, many questions remain, particularly with regard to the identity of predators, the episodic nature of selective predation events, and their apparent lack of lasting and significant impact on overall gene frequencies. We conclude that H. misippus presents both challenges and opportunities for studies on mimicry, and we suggest that linkage disequilibrium can be a useful generic indicator for Gestalt predation on polymorphic prey. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100, 180–194.

1. The aim of this paper is to investigate mechanisms of reinforcement between two semi-isolated semispecies of the African savannah butterfly Danaus chrysippus. The biogeography of colour genes suggests that four semispecies evolved in once isolated refugia. They expanded their ranges in response to Holocene climatic changes to form a hybrid zone in central-east Africa. 2. Danaus chrysippus is a superspecies within which cycles of alternating cladogenesis and reticulation among semispecies have probably operated over some 4 Myr. Semispecies are inter-fertile but show Haldane rule effects in crosses; gene flow is massive but subject to isolation by distance. 3. One semispecies shows linkage disequilibrium, vis- a-vis others, for haplotype, karyotype (W-linkage of colour genes which function as reproductive isolating barriers) and all-female broods caused by a male-killer endosymbiont. Introgression of colour genes between D. c. dorippus and D. c. chrysippus is constrained by sex linkage and male killing. 4. Reinforcement in hybrid zones comprises allochronic migration, assortative mating, (assumed) sex chromosome incompatibility and sex-ratio distortion. Gene introgression from D. c. dorippus to other semispecies is maintained by a high frequency of backcrossing between hybrid males and females of the latter.

SYNOPSIS The variation in cardenolide content found in 50 individuals of Danaus chrysippus is related to the type of food plant and the sequestering efficiency of the butterfly. The toxicity of the insects to bird predators is discussed in relation to the polymorphism found in species of Danaus.

This paper, which presents an annotated checklist of the ‘lower Nymphalidae’ (Libytheinae, Danainae, Satyrinae, Charaxinae), is the fourth in a series on the butterfly fauna of Mount Kilimanjaro. Four genera of lower Nymphalidae (Danaus, Amauris, Bicyclus, Charaxes), with a total of 11 included species, are known or believed to occur within the main forest zone, from c. 1800 to 2800 m. Of these, only three species of Charaxes (Charaxesberkeleyi, Charaxes ansorgei, Charaxes xiphares) may be restricted locally to this primary forest. The lower slopes fauna, below 1800 m, is considerably richer, with a total of 11 genera and 41 species listed (8 species of which extend into the forest zone). Possible additional species, dubious earlier records, problems with African subspecies of Danaus chrysippus, a need for more work on certain Satyrinae, and classification of the genus Charaxes are discussed. An identification key to the subfamilies of Nymphalidae, and the 19 genera of Libytheinae, Danainae, Satyrinae, Charaxinae that occur in Tanzania, together with a key to the adults of all the species of these four subfamilies considered to occur or have occurred on Kilimanjaro, with 206 colour images, are included as online Supplementary Information.

Mitochondrial and nuclear DNA information was analysed among four subspecies of the African Queen butterfly, Danaus (Anosia) chrysippus sensu lato (s.l.), along with four other Danaus species drawn from all three subgenera (D. (Danaus) plexippus, D. (Salatura) genutia, D. (A.) gilippus, D. (A.) eresimus) and two outgroup species from the same tribe, Tirumala septentrionis and Amauris niavius. A mitochondrial phylogeny derived from the 12S rRNA (347 bp) and COI (537 bp) loci indicates two very distinct haplotypes for subspecies D. (A.) c. dorippus, dorippus-1 and dorippus-2. Interestingly, dorippus-1, on the one hand, and all other D. (A.) chrysippus haplotypes, on the other, are the most distantly related clades within the genus and have different most recent ancestors from different subgenera, though sharing the common ancestor of the monophyletic genus. A phylogeny based on the EF1-α nuclear locus (400 bp) shows that the two well-separated mitochondrial lineages of dorippus are identical for this gene and reciprocally monophyletic to the other D. (A.) chrysippus lineages. Thus, nuclear and cytoplasmic phylogenies are not only discordant, but also suggest that both D. (A.) chrysippus s.l. and subspecies dorippus are polyphyletic. Paradoxically, four African subspecies, chrysippus-orange, chrysippus-brown, alcippus and dorippus, though substantially vicariant, hybridize extensively in East Africa wherever the ranges of two or more of them overlap. Linkage disequilibrium, and hence sexual isolation, in sympatry between colour (nuclear) genes and unlinked mitochondrial (cytoplasmic) loci is consistent across populations and therefore indicates the operation of positive natural selection. Together with data from previous experimental and field work, our results suggest that extensive hybridization occurs among once allopatric or parapatric lineages, that are now nascent species. We deduce that hybridism among lineages in sympatry is currently enforced, in the face of assortative mate choice, by a bacterial symbiont, Spiroplasma, a male-killer that forces females in female-biased populations to pair with heterotypic males. In discussion we emphasize that neither D. (A.) chrysippus s.l. as presently circumscribed, nor its component clades, conform to any established concept of species. © 2005 The Linnean Society of London.

Butterflies of the Danaidae family are considered to be toxic or distasteful due to the presence of cardiac glycosides sequestered from their larval food plants. Alcoholic extracts of specimens of Danaus chrysippus aegyptius and Amauris ochlea ochlea from southern Africa (Namibia, S.-Africa, Mozambique) were analyzed by thin-layer chromatography for these cardenolides. But only 4 of 75 specimens of D. chrysippus aegyptius contained trace amounts, all others including 13 specimens of A. ochlea ochlea were negative. Genetic analysis of the ouabain binding site of the Na(+), K(+)-ATPase revealed that both species do not present an amino acid replacement at the position 122, which otherwise makes the enzyme insensitive to cardenolides suggesting that other strategies of toxin tolerance must have been developed.

The plain Tiger Danaus chrysippus chrysippus (Linnaeus) (Insecta: Lepidoptera: Rhopalocera: Danaidae) is a widely distributed butterfly is rare in Andaman and Nicobar Islands. The scarcity of food plant Calotropis in the islands is the reason for rarity. The immature stages of the butterfly are discussed in the paper with some unusual observations such as laying of eggs in cluster and on the upper side of food plant, showing stress on survival of the fly.

—Molecular clocks based on sequence change in mitochondrial (mt) DNA have been useful for placing molecular phylogenies in their historical context, thereby enhancing evolutionary insight. Nonetheless, despite their importance to phylogeographers, the methodology is controversial. Here we report on two mitochondrial clocks for the butterfly genus Danaus based on sequences from the cytochrome c oxidase subunit I (COI) and small subunit 12S rRNA (12S) genes. Both clocks are, within the context of Danaus, reliable time-keepers, mutually consistent and, respectively, in agreement with a crustacean COI clock and a molluscan 12S clock. Though we have no fossils with which directly to calibrate sequence divergence rates for Danaus, the 12S molluscan and COI crustacean clocks chosen for comparison were calibrated to radiometrically dated geomorphological events. Our results indicate that the Danaus COI clock evolves approximately four times faster than the 12S clock. Differences between rates of sequence change in terminal sister-taxa are small and likelihood ratio tests do not reject a hypothesis that evolution has been clock-like. The species Danaus chrysippus is paraphyletic and, therefore, invalid. Danaus probably split from its sister-genus Tirumala around 4.9 ± 0.3 million years ago in the early Pliocene.

Subspecies chrysippus, dorippus and alcippus of the butterfly Danaus chrysippus differ at three biallelic colour gene loci. They have partially vicariant distributions, but their ranges overlap over a substantial part of central and East Africa, where hybridism is commonplace. We now report that the West African subspecies alcippus differs from other subspecies, not only in nuclear genotype but also in mitochondrial haplotype in both allopatry and sympatry. The maintenance of concordant nuclear and cytoplasmic genetic differences in sympatry, and in the face of hybridisation, is prima facie evidence for sexual isolation. Other evidence that suggests alcippus may be isolated from chrysippus and dorippus include differences in sex ratio (SR), heterozygote deficiency at one site and deduced differences in patterns of migration. We suggest that, within the hybrid zone, differential infection of subspecies by a male-killing Spiroplasma bacterium causes SR differences that restrict female choice, triggering rounds of heterotypic mating and consequent heterozygote excess that is largely confined to females. The absence of these phenomena from hybrid populations that test negative for Spiroplasma supports the hypothesis. The incomplete sexual isolation and partial vicariance of alcippus suggests that it is a nascent species.

A cross between queen butterflies of the Palaeotropical species Danaus chrysippus and the Neotropical D. gilippus was achieved with difficulty in both directions. Only one progeny (N=70) was reared comprising sterile males and inviable females in a precisely 1:1 ratio. Both prezygotic and postzygotic barriers to gene flow are strong. The result supports Haldane's Rule, to which we propose a minor amendment. The F1 hybrids were intermediate for background colour between the brown (genotype BB) of gilippus and orange (genotype bb) of chrysippus. Most F1 pattern characters were also intermediate. In polymorphic chrysippus populations, because Bb heterozygotes are brown, or nearly so, we suggest the B allele may have evolved towards dominance in sympatry. Hybrid males show positive heterosis for body size. The close similarity of male genitalia between the allopatric, genetically distant species chrysippus and gilippus, compared to their divergence between gilippus and its largely sympatric sister species eresimus, suggest that reinforcement of sexual isolation or reproductive character displacement have evolved in sympatry. © 2002 The Linnean Society of London.

Tujuan kegiatannya adalah : a) inventarisasi dan identifikasi jenis kupu-kupu di TWA Gua Pattunuang dan b) meningkatkan pemahaman murid Sekolah Dasar tentang peranan kupu-kupu di alam dan sumber keanekaragaman hayati. Manfaatnya adalah : kupu-kupu dapat menjadi sumber lapangan kerja dan pendapatan masyarakat di TWA Gua Pattunuang jika dilakukan secara berkelanjutan. Penelitian berbentuk survei dilaksanakan di TWA Gua Pattunuang dan pengenalan jenis kupu-kupu untuk murid SD 129 Inpres Bantimurung Kabupaten Maros, yang berlangsung pada bulan Maret sampai Mei 2017. Tahapan pelaksanaan kegiatan : 1) pemasangan transek dan pengamatan jenis kupu-kupu; 2) identifikasi kupu-kupu dan menghitung keanekaragamannya dengan Indeks Shannon-Wiener; dan 3) pengenalan jenis kupu-kupu ke murid Sekolah Dasar. Jenis kupu-kupu yang diperoleh sebanyak 20 individu yang terdiri dari 6 jenis famili Papilionidae (T. haliphron, T. helena, P. demoleus, P. demolion dan G. agamemnon) dan D. chrysippus dari famili Nymphalidae. Berdasarkan nilai Indeks Shannon-Wiener untuk semua jenis kupu-kupu mempunyai kriteria sangat rendah (H < 1). Saat kegiatan berlangsung, lokasi pengamatan berada dalam peralihan musim hujan ke musim kemarau. Kesimpulannya : 1) kupu-kupu yang ditemukan sebanyak 20 individu terdiri dari 6 jenis yang berasal dari famili Papilionidae dan Nymphalidae; 2) sebanyak 57.3% murid Kelas 5 SDN 129 Inpres Bantimurung telah mengenal jenis kupu-kupu dan 28.8% dari murid SD berprofesi sebagai penangkap kupu-kupu. Rekomendasi untuk kegiatan ini adalah kerjasama Universitas dan Pemda Kabupaten Maros untuk memperbanyak kegiatan sosialisasi manfaat kupu-kupu kepada murid sekolah yang bermukim di sekitar TWA Gua Pattunuang.Kata Kunci : kupu-kupu, Gua Pattunuang, Papilionidae, Nymphalidae

Large numbers of adults of certain species of butterfly flying in an apparently 'purposeful' manner are often noted by entomologists and the general public. Occasionally, these are recorded in the literature. Using these records we summarise information regarding the direction of movement in Australian butterflies and test whether there are consistent patterns that could account for known seasonal shifts in geographical range. The data were analysed using contingency tables and directionality statistics. Vanessa itea, Vanessa kershawi, Danaus plexippus, Danaus chrysippus and Badamia exclamationis flew predominately south in the spring-summer and north in the autumn-winter. Tirumala hamata has a strong southern component to its flight in spring but, as in Euploea core, appears non-directional in the autumn. For many supposedly known migratory species, the number of literature records are few, particularly in one season (mainly autumn). Thus, for Appias paulina, four of seven records were south in the spring-summer, as were six of nine records for Catopsilia pomona, and three of five for Zizina labradus. For Belenois java, flight records were only available for the spring and these showed geographical differences; predominantly north-west in northern Australia (Queensland) and south-west in southern Australia (Victoria, New South Wales). There were too few records for Papilio demoleus in the literature (four only) to draw any conclusions. Major exceptions to the seasonal trend of south in the spring and north in the autumn were Junonia villida, which showed a predominant north-westward direction in both seasons, and Eurema smilax, with a predominant southern or western flight in both seasons. We discuss these species specific trends in migration direction in relation to seasonal shifts in suitable habitat conditions, possible cues used in orientation and in timing changes in direction.

The aposematic savanna butterfly Danaus chrysippus seems to be an exception to the rule that Batesian models and Mullerian mimics are not polymorphic. Throughout Asia and in much of Africa the species is in fact monomorphic and polytypic, as expected. It is, however, polymorphic for 3–4 colour genes over a large area of central and eastern Africa, where deviant sex ratios and non-Mendelian segregations also occur. All-female broods are widespread and frequent, often outnumbering bisexual progenies and giving rise to heavily female-biased populations. Full and partial sex linkage, which is not controlled from the X or Y chromosomes, and male-biased broods also occur. Genetical analysis for the BC-autosome carrying colour genes suggests there are two, probably mitochondrial, cytotypes (microbe-induced early male death syndrome is considered unlikely) and an autosomal, incompatibility (I) gene, two alleles of which are male-specific killers. F2 and backcross matings by females heterozygous at the I-locus give progenies which are either thelygenic, all males dying at or soon after hatching, or bisexual but showing full or partial sex linkage. Male death is attributed to nuclear-cytoplasmic incompatibility (NCI). Females achieve reversion from a thelygenic to a bisexual line by mating with males of compatible (maternal) cytotype. A second NCI system causes meiotic drive in female oogenesis for a cytoplasmically compatible A-autosome, with no effect on SR. The array of non-Mendelian segregations for sex and two autosomes, with switching between bisexual and unisexual lineages, indicates autosomal intragenomic conflict with the outcome that one autosome is preserved from elimination in females and the other condemned in dead males. The geographical distribution of the colour marker genes suggests that the polymorphisms have a hybrid origin involving several once allopatric subspecies, some of which have divergent cytotypes. Secondary contact probably followed range expansion induced by the periodic spread of savanna during the Pleistocene and massive deforestation by man in recent times. The unusual extent of the hybrid zone is attributed to the large number of subspecies involved in its origin, and to introgression promoted by the butterfly's extensive migratory activity.

Non-Mendelian segregation involving three colour gene loci in the butterfly Danaus chrysippus is reported. In each of the four affected broods, two of the four phenotypes expected in the progeny were absent in both sexes. Operation of a system of balanced recessive lethals is ruled out by the low egg–adult mortality recorded. All gametes were produced in expected ratios and were viable. The results are interpreted to be consistent with an inherited prezygotic incompatibility mechanism which operates at the time of fertilization. Variable penetrance of the recessive a and c alleles in heterozygotes is significantly influenced, in the former case by sex, and in the latter by epistatic interactions with colour genes at the A and B loci. The combination of non-Mendelian segregation, variable incomplete dominance and epistatic interactions with other colour genes and sex supports the hypothesis, hitherto based largely on biogeographical evidence, that the polymorphic D. chrysippus population at Dar es Salaam, Tanzania may have a hybrid origin.

The diadem butterfly, Hypolimnas misippus, shows various anomalies in its presumed Batesian mimetic relationship with the distasteful danaine, Danaus chrysippus. If these anomalies are to be resolved and if reports of sexual selection in female diadems are to be correctly interpreted, an understanding of the genetics of the colour pattern polymorphism is essential. In an earlier study we developed a three-locus model (the A-M-S model) for colour pattern genetics in the diadem butterfly. Here, we show that the A locus segregates for body size as well as colour pattern. It is probably a supergene with at least four loci linked in coupling. Segregation for body size confirms several aspects of the A-M-S model: the ability of different genotypes to produce the same mimetic phenotype; the presence of a suppressor allele for hindwing white; and the effects of the A locus on the forewings of mm butterflies. Differences between the genetics of colour pattern in H. misippus and its model, D. chrysippus, favour a multiple origins rather than a selective maintenance hypothesis for the evolution of mimicry. Body size differences in both sexes must play a significant but as yet unexamined role in the maintenance of colour pattern polymorphism and in sexual selection in this species.

Samples of the polymorphic butterflyDanaus chrysippusare analysed from six well separated sites in East Africa. Morph-ratio clines are described for four diallelic genes A, B, C and L, each of which influences the visual phenotype. Each of the four clines has a different orientation, consistent with an hypothesis that the polymorphism originated from hybridization between a number of polytypic demes which have at various times undergone range expansion. Allopatric subspeciation in isolated Pleistocene refugia is postulated. The phenotype of each geographical race is shared with one of the morphs within the hybrid zone; other sympatrically maintained polymorphic forms are normally confined to the hybrid zone. Wright's isolation-by-distance model best explains the present distribution of gene frequencies. Morph-ratios differ significantly between the sexes and are sometimes associated with heterozygote excess; gametic and genotypic disequilibria are general throughout the region and suggest the clines are maintained by strong natural selection. Seasonal cycling of phenotype frequency is believed to result from extensive migratory movements rather than natural selection. Female-biased sex-ratio, which is also seasonal, and Haldane rule effects, result from hybrid breakdown when genetically distinct demes meet and interbreed. Oscillating sex-ratios and frequency of colour genes are functionally linked by negative feedback. The polymorphism owes its origin to allopatric evolution but is now maintained sympatrically.

Mullerian mimicry, in which there is convergence in coloration between unrelated unpalatable species, should lead to uniformity in appearance, not polymorphism, and so the occurrence in tropical Africa of unrelated species of unpalatable butterflies with corresponding polymorphic colour forms suggested a problem of special evolutionary interest. Field work in Uganda and Sierra Leone in 1964 72 demonstrated a statistical association between the occurrence and relative frequencies of polymorphic forms in Danaus chrysippus (Danainae) and Acraea encedon (Acraeinae) which was deemed as confirmation of a Mullerian relationship between them. There were, however, certain anomalies which at the time remained unresolved. Later, in 1976, it was discovered that what had been called A. encedon is in reality two sibling species. A. encedon and a new one, named as A. encedana. The two differ in the structure of both male and female genitalia and in the coloration and the food-plants of the larvae. The recognition of the additional species has enabled a re-assessment of the polymorphie Mullerian association with D. chrysippus. It emerges that, although there is a close qualitative and quantitative reciprocal mimetic relationship between A. encedana and D. chrysippus, the relationship between A. encedon and D. chrysippus is much weaker, and in places non-exitent. The possible origin of the mimetic polymorphism is discussed in terms of hybridization of previously allopatric and monomorphic populations which have met as a consequence of recent expansions of geographical range in all three species resulting from forest clearance and the spread of savanna-like conditions in previously forested areas.

The essential oil of the fruits of Periploca laevigata Aiton subsp. angustifolia (Labill.) Markgraf (Apocynaceae) from Lampedusa Island was obtained by hydrodistillation and its composition was analysed. The analyses allowed the identification and quantification of 64 volatile compounds belonging to different classes. The most abundant compounds were nonacosane, heptacosane, hentriacontane and δ-cadinene. Among the volatile compounds identified in the fruits of P. laevigata subsp. angustifolia, 31 are present in other taxa of Apocynaceae, 19 have antimicrobial activity and four are pheromones for the butterfly Danaus chrysippus. The possible ecological role of the volatile compounds found is briefly discussed.

The complete mitochondrial genome sequence of Danaus chrysippus (Lepidoptera: Nymphalidae: Danainae) was determined. The 15,236 bp long genome encodes 13 putative proteins, two ribosomal RNAs, 22 tRNAs and a non-coding AT-rich region. Its gene arrangement pattern is identical to most of other lepidopteran species. All protein-coding genes start with a typical ATN codon with the exception of COI gene which uses CGA as its initial codon; all PCGs terminate in the common stop TAA or TAG, except COI, COII, ND5 and ND4 which use single T as their stop codons. A total of 102 bp intergenic spacers and a total of 33 bp overlapping sequences are interspersed throughout the whole genome. The mitogenome harbors 22 txRNAs as those of most insect species and all tRNA genes evidence the typical clover leaf secondary structures with the exception of tRNAser (AGN) who loses its dihydrouridine (DHU) arm. The lrRNA and srRNA genes are 1339 and 783 bp, with the AT contents of 84.1 and 84.8%, respectively. The non-coding AT-rich region is 418 bp long, and contains the motif ATAGA followed by a 21-bp poly-T stretch and a microsatellite-like (AT)9 element preceded by the ATTTA motif.

Subahar TSS, Yuliana A (2010) Butterfly diversity as a data base for the development plan of Butterfly Garden at Bosscha Observatory, Lembang, West Java. Biodiversitas 11: 24-28. Change of land use and the increasing number of visitors to Bosscha area was one factor for the development plan of butterfly garden in the area. The objectives of this research were to examine butterfly diversity and its potential for development plan of butterfly garden. Butterfly diversity and its richness conducted by standard walk methods. Host plantand larval food plant was recorded during butterfly survey. Public perception on the development plan of butterfly garden was examined by questionnaire. The results showed that 26 species of butterfly was found in Bosscha area and Delias belisama belisama was the most dominant species. Public perceptions consider that the development plan of butterfly garden will give benefit to the community; not only providing new insight (40.41%), additional tourism object (23.97%) and will gave aesthetical value (17.12%). Twelve local speciesshould be considered for development plan of butterfly garden: Papilio agamemnon, P. demoleus, P. memnon, P. sarpedon, Delias belisama, Eurema hecabe, Danaus chrysippus, Argynnis hyperbius, Cethosia penthesilea, Hypolimnas misippus, Melanitis phedima and Euthalia adonia. Host plant: Bougainvillea spectabilis, Citrus aurantium, Lantana camara, Macaranga tanarius and food plants: Citrus aurantium, Cosmos caudatus, Eupatorium inulifolium, Gomphrena globosa, Hibiscus rosa-sinensis, Lantana camara, and Tithonia diversifolia.Key words: host plant, butterfly diversity, Bosscha Observatory, standard walk.

The Batesian mimetic swallowtail butterfly Papilio dardanus exhibits numerous distinct wing colour morphs whose evolutionary origins require large phenotypic shifts. A phylogenetic framework to study the history of these morphs was established by DNA sequencing of representative subspecies from sub-Saharan Africa and Indian Ocean islands. Two mitochondrial genes and the nuclear internal transcribed spacer marker revealed deeply separated eastern and western African mainland lineages, plus one lineage each on Madagascar and Grande Comore. These markers showed very little polymorphism within lineages. In contrast, markers genetically linked to the mimicry locus H, including the transcription factor invected and two adjacent amplified fragment length polymorphisms-derived sequences, showed high nucleotide diversity but were not geographically structured. Variation in the unlinked wingless gene showed a similar pattern, rejecting the hypothesis that high level of variation in the H region is due to balancing selection exerted by the phenotypes. The separation from a common ancestor with Papilio phorcas estimated at 2.9 Ma coincides with the origin of a mimicry model, Danaus chrysippus. However, the model reached Africa only at the time of the internal splits of P. dardanus mtDNA groups, here estimated at 0.55-0.94 Ma. The nuclear genome shows less geographic structure and may not track recent population differentiation, suggesting that widespread mimicry morphs have arisen early in the evolution of the P. dardanus lineage, although after the male-female dimorphism which is ancestral. The current wide distribution of P. dardanus and population subdivision evident from mtDNA may have been achieved only with the spread of the models across Africa.

Summary In some populations of the African butterfly, Acraea encedon, there are two kinds of females, one producing offspring in a normal 1:1 sex ratio, the other producing females only; in other populations the sex ratio is apparently normal. All-female broods had hitherto been mainly associated with populations in which field sampling revealed an excess of females. The all-female brood trait is described from a population at Dar es Salaam which field sampling suggested was normal, and this indicates that the trait may be much more widespread and common than had previously been supposed. This discovery also extends the known distribution of the trait across Africa from Sierra Leone to eastern Tanzania. The butterfly is also a polymorphic Mullerian mimic of Danaus chrysippus, which is a highly unusual phenomenon as Mullerian mimicry is almost invariably monomorphic. The relative frequencies of two corresponding colour forms of the two species of butterflies at Dar es Salaam adds support to the hypothesis that they are indeed Mullerian mimics. The results of breeding experiments suggest that the polymorphic forms in Acraea encedon are allelic with dominance.

Female-biased sex ratios in Danaus chrysippus (Linnaeus) (family Nymphalidae, subfamily Danainae, tribe Danaini) populations are attributed to the action of an endosymbiotic 'male-killer' bacterium of the genus Spiroplasma. In stark contrast to the extensive geographic range of their host, the Spiroplasma appears to be restricted to East Africa, where four African D. chrysippus subspecies exist sympatrically and form a hybrid zone. In this study, specimens collected at three sample sites within the hybrid zone were screened for Spiroplasma infection. The findings demonstrate that, within the hybrid zone, the frequency of Spiroplasma infection varies both spatially and temporally. Host genotype at three biallelic wing pattern loci, representative of subspecific genetic divergence, is correlated to Spiroplasma infection. Linkage between the frequency of the recessive a allele and Spiroplasma is established, suggesting the presence of a linked allele controlling infection susceptibility. In addition, a negative correlation between D. chrysippus forewing length and Spiroplasma infection is identified, suggesting that infection has a deleterious effect on body size. © 2007 ICIPE.

Anecdotal evidence has long suggested that the African queen butterfly, Danaus chrysippus (L.), is migratory. More recently, records of rapid and seasonal changes in the frequencies of different colour pattern morphs in Tanzania, Ghana, and Kenya have provided further evidence. This paper demonstrates similarly rapid changes in the frequencies of different mitochondrial haplotypes in Nairobi, Kenya over a two-week period. These changes are correlated with changes in the frequencies of homozygous and hybrid colour forms, and are consistent with colour pattern/haplotype associations that have been observed on a continental scale. We conclude that they can only be explained by migration and that mtDNA analysis can be usefully and generally applied in studies of migratory movement.Journal of East African Natural History Vol. 92 (1&2) 2003: pp. 119-125

On August 15, 2004, three specimens of Danaus chrysippus (Linnaeus, 1758) were caught at the mouth of the Neretva River near the town of Ploče in South Dalmatia. This is the first record of this migratory species in Croatia. A review of its zoogeographical characteristics and distribution in Europe is given. The results have been discussed and further study of this species is recommended., Tri primjerka vrste Danaus chrysippus (Linnaeus, 1758) ulovljena su 15. kolovoza 2004. godine na ušću rijeke Neretve, blizu grada Ploča u južnoj Dalmaciji. To je prvi nalaz ove migratorne vrste za Hrvatsku. Prikazane su zoogeografske značajke i rasprostranjenost spomenute vrste u Europi. Rezultati su raspravljeni i preporučeno je daljnje istraživanje ove vrste.

On August 15, 2004, three specimens of Danaus chrysippus (Linnaeus, 1758) were caught at the mouth of the Neretva River near the town of Ploce in South Dalmatia. This is the first record of this migratory species in Croatia. A review of its zoogeographical characteristics and distribution in Europe is given. The results have been discussed and further study of this species is recommended.

Total and differential haemocyte counts following starvation and feeding resumption, ventral nerve cord transection, and brain implantation followed by methoprene application and 20-hydroxyecdysone injection were studied in Danaus chrysippus (Linnaeus) larvae. The results revealed that initial starvation and transection of the ventral nerve cord before the fifth abdominal ganglion in fifth instars of D. chrysippus led to a reduced total haemocyte count, on the one hand, and to a higher variation in percentage of different haemocyte types, on the other. While feeding resumption, brain implantation and 20-hydroxyecdysone treatments reversed the effect of starvation and nerve cord transection, methoprene application produced a decline in total number of haemolymph cells.

The lesser wanderer butterfly, Danaus (Anosia) chrysippus petilia (Stoll) (Lepidoptera: Danainae), has been treated as a subspecies for the last 100 years. New mitochondrial DNA sequence data for D. petilia, in conjunction with allozyme, structural, morphometric and pattern characters, constitute a compelling case for its specific rank. The holotype of D. petilia has never been located and, as the type location is uncertain, a neotype is designated. Fresh material was collected and examined for this project. Danaus petilia and D. chrysippus have been separated at Lydekker's Line for an estimated 1.1 million years, and they remain interfertile. However, as an allopatric taxon, diagnosable from D. chrysippus, D. petilia merits specific status under the phylogenetic species concept. The following pairs of D. chrysippus subspecies are considered to be synonyms, the first member of each pair having priority: chrysippus + aegyptius (subspecies), klugii + infumata (hybrid phenotypes) and orientis +liboria (subspecies).

The synthesis of (E)-3,7-dimethyl-2-octene-1,8-diol (1), which was isolated from the hairpencils of male Danaus chrysippus (African Monarch), was investigated. The key step of the sequence involves asymmetric desymmetrization of the 1,3-propanediol 7 with lipase, in which high enantioselectivity was observed. Total synthesis afforded (S)-1 in 12 steps and 26% overall yield from readily available geraniol.

Many insects carry maternally inherited bacteria which kill male offspring. Such bacteria will spread if male death benefits the female siblings who transmit the bacterium, and they are therefore expected in insects with antagonistic sibling interactions. We report that the butterfly Danaus chrysippus is host to a maternally inherited male-killing bacterium. Using diagnostic PCR and rDNA sequence, the bacterium was identified as a Spiroplasma closely related to 2 ladybird beetle male-killers and the tick symbiont Spiroplasma ixodetis. The male-killer was found to have a geographically restricted distribution, with up to 40% of females being infected in East Africa, but no detectable infection in small samples from other populations. Danaus chrysippus is a surprising host for a male-killer as its eggs are laid singly. This suggests that the ecological conditions permitting male-killers to invade may be more widespread than previously realized.

Female-biased sex ratios occur in many insect species as a consequence of infection by maternally-inherited male-killing bacterial endosymbionts. In this paper, we revise the research conducted on the phenomenon of male-killing in African nymphalid butterflies, with special focus on the cases of Danaus chrysippus , Acraea encedon and Acraea encedana . The evolution of male-killing in each case was addressed, together with the phylogeny of male-killers that were identified from this group. Moreover, the potential impacts that male-killers might impose on the evolution of their butterfly hosts were thoroughly investigated. In the end of this review, we present a number of unanswered questions to be targeted by future research work on the male-killing in these butterflies.

There are many good general reviews of the evolution of mimicry1xMimicry in Plants and Animals. Wickler, W. See all References, 2xTurner, J.R.G. : 141–161See all References, including Edmunds's own excellent book3xDefence in Animals. Edmunds, M. See all References3. Our perspective dealt only with the paradox that mimicry rampantly diversifies in morphs, subspecies and species, even though simple theory explains only convergence in a reduction of diversity. (Also, we did refer to MacDougall and Dawkins, see Ref. 31 in our original article; however, their idea is now criticized4xSee all References4.) We concentrated on butterflies because most studies relating theory, field data and genetics are done in this group.We very much doubt the implication that butterflies are special; mimicry ought to work the same in less well known organisms. Edmunds and Golding suggest that female sex-chromosome heterogamety and female-limited mimicry in butterflies are related. However, mimetic sexual dimorphism in butterflies is due, in part, to sexual selection acting on males5xThe Descent of Man, and Selection in Relation to Sex (2nd edn). Darwin, C. See all References, 6xTurner, J.R.G. Biol. J. Linn. Soc. 1978; 10: 385–432CrossrefSee all References, 7xVane-Wright, R.I. : 251–253See all References, and similar dimorphisms are also common in male-heterogametic organisms, such as mammals. In only one case – Papilio glaucus in N. America8xHagen, R.H. and Scriber, J.M. J. Hered. 1989; 80: 179–195See all References8 – is a mimetic-switch gene linked to the female-specific W chromosome; most mimetic polymorphisms are inherited autosomally, with sex chromosomes merely serving to switch between mimetic and nonmimetic patterns through their effect on sex. Although no cases of female-limited mimicry are known in organisms other than butterflies, we don't see a good reason for this: where ecologically feasible, there are cases of male-limited mimicry in Lepidoptera9xVane-Wright, R.I. J. Zool. 1975; 177: 329–337CrossrefSee all References9, for instance in diurnal male moths that search for females that rest by day.The assertion that butterflies have become diurnal because of unpalatability is clearly wrong. Basal groups like Hesperiidae are chiefly palatable, and the palatability of most Dismorphiinae, Coliadinae, Lycaenidae and Libytheinae argues for unpalatability evolving within the Pieridae and Nymphalidae, rather than before their radiation. Mimicry is associated with diurnality in Lepidoptera; but this is hardly surprising given that nocturnal predators rarely use details of colour and pattern when hunting prey. Rapid and unpredictable flight (Castniidae, Hesperiidae and Charaxinae), low profitability (Satyrinae and Lycaenidae), and camouflaged undersides (many Nymphalidae) are among other common attributes of Lepidoptera that allow diurnality. Unpalatability, warning colour and mimicry are clearly among adaptations that permit diurnality, but they are neither required nor unique to butterflies. Anyone who has been stung by a Paraponera and swarmed over by Crematogaster will be able to inform about great differences in the unpleasantness of ants, and it seems probable that similar palatability spectra exist in spiders and Diptera. Mimetic butterflies are better studied, rather than having special kinds of palatability or mimicry.Gordon and Smith resent our correct citation of their group's earlier ideas on Batesian overload in Danaus. However, we also reported (p. 463 in our original article) their current hypothesis: ‘recent contact… of previously isolated geographical races has been suggested as a nonadaptive cause of polymorphism in these species’. Although we did not say so in the perspective, secondary contact is not a good deus ex machina for explaining mimetic polymorphism. It has recently been found that in both Acraea encedon10xSee all References10 and Danaus chrysippus (F. Jiggins, pers. commun.) sex ratio distortion is a result of infection by male-killing bacteria rather than hybrid breakdown. We have no doubt that similar nonhistorical ideas will explain other puzzling features of the genetics of Danaus and Acraea. In any case, geographic isolation in a refuge cannot by itself cause an initially unfavourable colour pattern to spread; drift and/or fluctuating selection are required initially, and these could also take place during parapatric contact, as in Heliconius11xMallet, J. and Turner, J.R.G. : 262–280See all References11, making allopatry unparsimonious. The Danaus anomaly also stubbornly refuses to disappear via an accusation of beanbag genetics. We must still explain an area larger than Europe with unexpected mimetic polymorphism. We suggest that this and other Mullerian polymorphisms might be maintained by spatiotemporal heterogeneity, as described in our article. Mimetic selection in Danaus may be fairly weak in the area of polymorphism, and balanced by fairly strong gene flow (large-scale migrations), leading to a wide band of polymorphism between areas of stable relative purity. Similar patterns of current selection are a likely source of widespread polymorphism in Mullerian mimetic ladybirds12xMelanism. Majerus, M.E.N. See all References12.

Danaus chrysippus is a polytypic tropical butterfly which is divided in Africa into five geographical races, each characterised by colour genes which are identifiable in the phenotype. The genetics of the races is known except for liboria which is described here for the first time. The origin of the races is assumed to be allopatric but expansion into East Africa since the Pleistocene has produced an extensive polymorphic hybrid zone. Regular scoring of phenotype frequencies at Dar es Salaam over two years suggests that replicated seasonal oscillations in gene frequency reflect migratory activity rather than natural selection in the two races (dorippus and liboria) studied. The hypothesis is supported by direct observations of north-south return migration within East Africa and numerous reports of migratory activity elsewhere. Migration is related to the cyclical north-south movement of the inter-tropical convergence zone and consequent changes in wind direction, rainfall and temperature.

The African butterfly Danaus chrysippus, like other members of the family Danaidae, feeds as a larva on species of milkweeds (Asclepiadaceae). It has been demonstrated in a sample from a West African population that only a minority of adult D. chrysippus accumulated detectable amounts of poisonous and presumably emetic cardenolides from their larval foodplants. It has also been shown in D. chrysippus and the related monarch butterfly (D. plexippus) that the cardenolide content of adults varies with that of their respective milkweed foodplants. Three population samples of D. chrysippus from sites in East Africa are analysed for cardenolides using the spectrophotometric technique refined by Brower, Edmunds & Moffitt (1975), which allows the assay of individual butterflies. The East African samples all have a palatability spectrum with a considerably higher percentage of cardenolide-containing individuals than the West African one. This is discussed in terms of its effect on potential bird predators and the ‘automimicry’ of emetic by palatable individuals. In East Africa D. chrysippus is polymorphic and appears to act as a model in a complex mimicry ring, whereas in West Africa it is monomorphic and may have ‘shed’ most of its mimics. This latter hypothesis is discussed in the light of our results.

Males ofDanaus butterflies possess two binate glandular organs: abdominal hairpencils, which release aphrodisiac pheromones during courtship behavior, and pockets on the hindwings. Between these two types of organs contacts are established behaviorally: the hairpencils are dipped into the alar pockets (Fig. 2). GLC analyses of hairpencil and wing pocket extracts fromDanaus chrysippus show that this contact behavior is a prerequisite for the synthesis of the ketonic pheromone component (for which the term “danaidone” is suggested here) in physiologically normal amounts. Danaidone occurs on the hairpencils only i) after the males have ingested pyrrolizidine alkaloids as precursors, and ii) after the hairpencils have been dipped into the wing pockets. The appearance of danaidone in the wing pockets also depends on the ingestion of alkaloidal precursor, but is not affected by the above mentioned contact behavior. Mechanisms by which contact behavior might control pheromone biosynthesis are discussed.

When ration (100%, 50% and 25%) and feeding duration (3, 6, 9, 12 and 24 hr/day) were restricted Danaus chrysippus extended larval period from 6 to 18 days; however, the pupal duration remained constant (6.3 ± 0.4 days) in all the groups. Those fed less than 50% ration and 6 hr/day suffered heavy mortality. Restriction of feeding duration is partially compensated by increasing feeding rate. Despite these restrictions of feeding schedule, the larvae in either series exhibited equal efficiencies of assimilation (44%), and conversion (40%). Hence, the restriction resulted in the production of miniature pupae, whose efficiency to retain body substances in the adult is low (

Danaus chrysippusis a danaid butterfly commonly found throughout Africa. It feeds, in its larval stage, on species of milkweed (fam. Asclepiadacae). If the milkweed species has a high cardenolide content, then the larva as well as its pupa and the subsequent adult will be toxic to birds. If, however, the larva has fed on a species of milkweed with little or no cardenolide content, then it will be palatable to birds and so will the pupa and the adult. All the three stages (larval, pupa and adult), whether palatable or not, are subject to attack by birds. The larvae, on the other hand, also act as hosts to a large number of parasitoids. In this paper a theoretical model is described which incorporates the following three assumptions that have been verified experimentally: (i) female butterflies prefer to oviposition the species of plants on which they had developed; (ii) birds are able to distinguish between externally identical larvae from the shape of the plant on which the larvae are feeding; (iii) female parasitoids select, for oviposition,D. chrysippuslarvae feeding on plants toxic to vertebrate predators. The simulations using the theoretical model predicts that parasitoids will prefer to lay their eggs in toxic larvae that are less likely to be predated by birds. This predation makes it advantageous for larvae to feed on non-toxic plants. This larval advantage is countered at the adult stage by bird predation. As one type of butterfly becomes too common, the other type will be at an advantage. The polymorphism for palatability is maintained primarily through selection at the larval stage and not by bird predation on the adult butterflies. The experimental data supporting the above assumptions are discussed. Computer simulation predicts the behaviour of birds and parasitoids as it affects the relative frequency of toxic and non-toxic butterflies.

Danaus chrysippus was reared in the laboratory from stock obtained from Kampala (Uganda), Nairobi (Kenya), Dar-es-Salaam (Tanzania) and Freetown (Sierra Leone) and wild-caught samples from Nigeria, southwest Africa and Tanzania were also analysed. D. plexippus was reared on the same plants for comparison. It was found that the adult D. chrysippus is a poor and inconsistent storer of cardiac glycosides compared with D. plexippus , and contained, chiefly, highly polar cardenolides. Populations in East Africa are, on the whole, more efficient storers than those from West Africa, a factor which may contribute to the dearth of mimics in West Africa. Furthermore there appears to be a genetic element in the storage capacity of these butterflies, not merely a ‘mirror’ effect, depending on the cardenolide content of their food plants. Differences in storage capacity were shown between morphs alcippus from both Sierra Leone and Dar-es-Salaam and aegyptius from Tanzania and Dar-es-Salaam, reared side by side on the same tested food plants. In both cases aegyptius was the better storer but in other broods from Kenya, reared on Asclepias rich in cardenolides, this morph was negative for these substances. During the investigation strains of Asclepias curassavica were found which contained calotropin, but lacked calactin. D. plexippus reared on these plants also lacked calactin, but sequestered and stored it when fed on Gomphocarpus fruticosus which contained both substances. The methods for analysing cardenolides of this type are described. Maps are presented showing the distribution of the three principal morphs of D. chrysippus and the form albinus in Africa.

The sudden weight loss after eclosion in the butterflies Acraea horta, Danaus chrysippus and Papilio demodocus is largely due to diuresis. The potassium concentration of the haemolymph is approximately halved as a result, and extensive diuresis leads to increases in sodium concentration and total osmolarity. The isolated Malpighian tubules of all three species are stimulated to fast rates of fluid secretion by cyclic AMP and by homogenates of the brains and corpora cardiaca. The tubules of Papilio lose their sensitivity to stimulation after the first day of adult life, and ingestion of a large volume of artificial nectar by this butterfly does not cause diuresis.