Papilio Memnon Research Articles

Complete mitochondrial genome data and phylogenetic analysis of Papilio macilentus Janson, 1877 (Lepidoptera: Papilionoidea: Papilionidae)

In the present study, the complete mitochondrial genome (mitogenome) of the Papilio macilentus (Lepidoptera: Papilionoidea: Papilionidae) was sequenced by next-generation sequencing method. The mitochondrial genome is a circular DNA molecule of 15,264 bp in size with 80.7% AT content, including 37 genes (13 protein-coding genes, 2 rRNA genes, and 22 tRNA genes), and a long non-coding region (Control region). All protein-coding genes are initiated by ATN codons, and terminated with TAA, TAG, or single T. All tRNAs can be folded into common clover leaf secondary structure, except trn-S1. Phylogenetic analyses based on 13 protein-coding genes and 2 rRNA genes using maximum likelihood and Bayesian inference confirmed that P. macilentus and Papilio memnon are clustered into a clade, and revealed the relationships between Papilionini, Troidini, Teinopaippini and Leptocircini.

Diversity and the role of butterflies species at PT Permata Sawit Mandiri, West Kalimantan

The use of chemicals indicates a cost inefficiency and makes oil palm plantations considered environmentally unfriendly causing a decrease in environmental quality. These problems indicate the need to increase the role of biodiversity to optimize oil palm productivity and be environmentally friendly. Based on this, research is needed on the diversity and role of butterfly species in palm oil agrosystems. The purpose of this study is to identify the species of butterflies in various land cover and analyzes the role of butterflies in the palm oil agrosystem. The research was conducted at PT Permata Sawit Mandiri, West Kalimantan. Diversity data was obtained through pollard transect and trapping methods, while role data was obtained through the literature study method. The pollard transect method was carried out along a line measuring 2 × 1,000 m long and 10 m wide at 08.00-11.00. The trapping method was carried out by placing 5 traps in each observation line for 3 × 24 hours. The results showed that there were 190 individuals belonging to Papilionidae (3 species), Pieridae (8 species), Nymphalidae (37 species), Lycaenidae (7 species), and Hesperidae (4 species). There are variations of butterfly species between land covers with the highest number of species found in rubber gardens (27 species). 7 species of butterflies have the potential as oil palm pollinators with the highest potential owned by Elymnias hypermnestra. The Papilionidae (Graphium agamemnon, Papilio polytes, Papilio memnon) can be environmental bioindicators because more sensitive to the environment.

Variations in species diversity and the role of butterflies in oil palm agroecosystems at PT BSM South Sumatra

Reducing the use of chemical fertilizers and controlling pests/chemical diseases is one of the prerequisites for the realization of sustainable oil palm plantation management. In line with that, the paradigm of green economic development requires optimizing the utilization of biodiversity in various aspects of managing oil palm plantations in Indonesia. Therefore, research is needed that can describe the variation and role of biodiversity (one of which is butterfly taxa) in the agro-ecosystem of oil palm plantations. This study aims to identify variations in species diversity and the role of butterflies in the agro-ecosystem of oil palm plantations. Observation of butterfly species diversity was carried out on seven types of oil palm plantation land cover at PT BSM. The method used is pollard transect (2 km with a width of 10 m) and trapping. The role of various types of butterflies is carried out through a literature search. The results showed that there were 163 individuals from 30 species and consisted of 4 families namely Papilionidae, Pieridae and Lycaenidae. The highest diversity and richness index values were found in smallholder plantations 1 (H′=2.42) and (Dmg=4.01), while the lowest were found in young oil palm areas (H′=0.87) and (Dmg=0. 99). The highest evenness index value was found in shrubs (E=1) while the lowest was in young oil palm land cover (E=0.54). Butterfly species that have potential as environmental bioindicators are Tanaecia pelea, Neptis hylas, Cupha erymantis, Neptis ida, Danaus affinis, Junonia atlites, Leptosia nina, Catopsilia pyranthe, Rapala iarbus, Papilio memnon, Losaria coon, Papilio polytes, Spindiasis lohita and Eurema hecabe. While the pollinators are Ypthima baldus, Catopsilia pyranthe, Hypolimnas bolina, Papilio polytes, and Neptis hylas.

Keanekaragaman Jenis Kupu-Kupu Di Air Terjun Kedung Kandang Nglanggeran

Penelitian ini bertujuan untuk mengetahui keanekaragaman jenis kupu-kupu di kawasan Air Terjun Kedung Kandang. Jenis penelitian yang dilakukan adalah deskriptif-eksploratif dengan pengambilan data menggunakan metode jelajah atau cruising method secara luring yang dilakukan pada bulan Maret-Agustus 2021. Waktu pengambilan data dilakukan pukul 08.00-12.00 WIB dilanjutkan pukul 14.00-16.00 WIB dan berlokasi di Air Terjun Kedung Kandang Nglanggeran, Kabupaten Gunungkidul, Daerah Istimewa Yogyakarta. Berdasarkan data pengamatan diperoleh hasil keberadaan Nymphalidae sebesar 59%, Pieridae 20%, Papilionidae 12%, dan Lycanidae 9%. Jenis kupu-kupu yang melimpah yaitu Papilio memnon dari famili Papilionidae. Mycalesis sp. dari famili Nymphalidae, Eurema sp. dari famili Pieridae, dan Jamides alecto dari famili Lycanidae. Sementara, hasil nilai H’ menunjukan angka 0,34. Nilai kemerataann jenis menunjukan nilai 0,113. Nilai tersebut mendekati 0 maka sebaran individu antar spesies tidak merata. Indeks dominansi sebesar 0,126 atau mendekati 0 maka tidak ada spesies yang mendominasi di kawasan Air Terjun Kedung Kandang. Sedangkan indeks kekayaan jenis menunjukan nilai 3,63 atau tergolong sedang. Berdasarkan penelitian yang telah dilakukan, dapat ditarik kesimpulan bahwa Keanekaragaman Jenis Kupu-Kupu Di Air Terjun Kedung Kandang tergolong kecil/rendah.

Identifikasi Jenis Kupu-kupu (Lepidoptera) di Taman Wisata Alam Baning Kabupaten Sintang

Baning Nature Park is located on the Ana River, Sintang District, Sintang District, West Kalimantan Province, is an ex-situ conservation area of ​​dry lowland plants with various collections of plant habitus, both from trees, shrubs, herbs, lianas, and others. other. The existence of butterflies in a habitat has a role as pollinating insects which have a very important position in maintaining the balance of nature. Butterflies aside from being natural pollinating agents, can be used as an indicator in testing environmental quality, by observing the state of their habitat. This study aims to find out what types of butterflies are found in the Baning Nature Park and their ecological roles. This research is a type of exploratory research. Data collection was carried out using a belt transect, by circling the Baning Nature Park. Catching butterflies is done using sweeping technique. Observations made at the Baning Nature Park found 15 species in 4 different families. These species namely; Arhopala centaurus, Leptosia nina, Appias olferna, Eurema blanda, Catopsilia pomona, Eurema hecabe, Junonia hedonia, Junonia atlites, Junonia iphita, Hypolimnas bolina, Phaedyma columella, Elymnias hypermnestra, Tanaecia trigerta, Papilio memnon, and Papilio polytes. The ecological role of the butterfly is as a pollinating organism. Nectar-producing plants in Baning Nature Park which are frequently visited by butterflies, namely, the species Psydrax dicoccos and Ixora sp.

Swallowtail butterflies and hawkmoths contribute equally to the pollination of Habenaria dentata (Orchidaceae)

Mixed pollination system with both diurnal and nocturnal visitors contributing to pollen transfer and seed set have been documented in many angiosperm species. The flowers of Habenaria species are usually green or white, with a long nectar secreting spur, and are usually interpreted as specialized for pollination by nocturnal moths. With a few exceptions, nocturnal moths and diurnal butterflies are both reported as pollinators of Habenaria species. However, the relative contributions made by moths versus butterflies to the reproductive success in the same orchid species has rarely been studied. We investigated the floral morphology, pollinators, pollination efficiency, and breeding system of Habenaria dentata in southern China. We showed that two species of diurnal swallowtail butterflies (Papilio helenus and Papilio memnon) and a species of crepuscular hawkmoth (Eupanacra mydon) were this orchid's effective pollinators. The pollinaria were attached to the eyes of these three insect species. Pollinator exclusion experiments showed that pollinarium removal and pollinium deposition rates on stigmas by butterflies (47.51% and 38.64%, respectively) were not significantly different from those made by hawkmoths (50.70% and 34.47%, respectively), indicating that diurnal and crepuscular visits contributed equally to reproductive success. As natural rates of insect-mediated pollination of our population of H. dentata resulted in fruit set rates of 93.33%, rejected the prospect that this population was pollinator limited. Hand pollination experiments indicated that H. dentata was self-compatible but without autonomous self-pollination. The proportion of well-developed seeds with large embryos in self-pollinated fruits was significantly lower than in cross-pollinated fruits, suggesting natural selection favors xenogamy. Our results highlighted the importance of both butterflies and hawkmoths contributing to the reproductive success of a rare orchid with a mixed pollination system.

Molecular phylogeny of swallowtail butterflies (Lepidoptera: Papilionidae) based on mitochondrial cytochrome c oxidase I (COI) gene from Bangladesh

Swallowtail butterflies (Papilionidae) are among the best-known insects in the world. Mitochondrial cytochrome c oxidase I (COI, 642 bp) gene from nine species under four genera of swallowtail butterflies was sequenced and submitted to the GenBank. Basic local alignment search tool (BLAST) analysis showed that the COI gene sequences of these nine butterflies matched those of respective species from different geographical areas (99-96% similarity). The interspecific genetic divergence among swallowtail species was between 0.01–0.15%. Phylogenetic tree constructed using the maximum likelihood algorithm showed that the species of swallowtail butterflies were originated from a common ancestor where Papilio polytes, Papilio nephelus, Papilio demoleus, Papilio memnon and Papilio polymnestor belonged to a cluster. On the other hand, Graphium agamemnon, Atrophaneura varuna and Pachliopta aristolochiae belonged to another cluster. In the TCS haplotype network, considerable genetic divergence was observed among the nine species of swallowtail butterflies. Both phylogenetic and haplotype analyses showed that Graphium doson was genetically most divergent among the swallowtail butterflies studied in the present study. These results indicate that Graphium spp. might be the good model system to uncover many biological processes like color vision and image processing in organisms.

Doublesex Controls Both Hindwing and Abdominal Mimicry Traits in the Female-Limited Batesian Mimicry of Papilio memnon.

Papilio butterflies are known to possess female-limited Batesian mimicry polymorphisms. In Papilio memnon, females have mimetic and non-mimetic forms, whereas males are monomorphic and non-mimetic. Mimetic females are characterized by color patterns and tails in the hindwing and yellow abdomens. Recently, an analysis of whole-genome sequences has shown that an approximately 160 kb region of chromosome 25 is responsible for mimicry and has high diversity between mimetic (A) and non-mimetic (a) alleles (highly diversified region: HDR). The HDR includes three genes, UXT, doublesex (dsx), and Nach-like, but the functions of these genes are unknown. Here, we investigated the function of dsx, a gene involved in sexual differentiation, which is expected to be functionally important for hindwing and abdominal mimetic traits in P. memnon. Expression analysis by reverse transcription quantitative PCR (RT-qPCR) and RNA sequencing showed that mimetic dsx (dsx-A) was highly expressed in the hindwings in the early pupal stage. In the abdomen, both dsx-A and dsx-a were highly expressed during the early pupal stage. When dsx was knocked down using small interfering RNAs (siRNAs) designed in the common region of dsx-A and dsx-a, a male-like pattern appeared on the hindwings of mimetic and non-mimetic females. Similarly, when dsx was knocked down in the abdomen, the yellow scales characteristic of mimetic females changed to black. Furthermore, when dsx-a was specifically knocked down, the color pattern of the hindwings changed, as in the case of dsx knockdown in non-mimetic females but not mimetic females. These results suggest that dsx-a is involved in color pattern formation on the hindwings of non-mimetic females, whereas dsx-A is involved in hindwing and abdominal mimetic traits. dsx was involved in abdominal and hindwing mimetic traits, but dsx expression patterns in the hindwing and abdomen were different, suggesting that different regulatory mechanisms may exist. Our study is the first to show that the same gene (dsx) regulates both the hindwing and abdominal mimetic traits. This is the first functional analysis of abdominal mimicry in butterflies.

Genomic architecture and functional unit of mimicry supergene in female limited Batesian mimic Papilio butterflies.

It has long been suggested that dimorphic female-limited Batesian mimicry of two closely related Papilio butterflies, Papilio memnon and Papilio polytes, is controlled by supergenes. Whole-genome sequencing, genome-wide association studies and functional analyses have recently identified mimicry supergenes, including the doublesex (dsx) gene. Although supergenes of both the species are composed of highly divergent regions between mimetic and non-mimetic alleles and are located at the same chromosomal locus, they show critical differences in genomic architecture, particularly with or without an inversion: P. polytes has an inversion, but P. memnon does not. This review introduces and compares the detailed genomic structure of mimicry supergenes in two Papilio species, including gene composition, repetitive sequence composition, breakpoint/boundary site structure, chromosomal inversion and linkage disequilibrium. Expression patterns and functional analyses of the respective genes within or flanking the supergene suggest that dsx and other genes are involved in mimetic traits. In addition, structural comparison of the corresponding region for the mimicry supergene among further Papilio species suggests three scenarios for the evolution of the mimicry supergene between the two Papilio species. The structural features revealed in the Papilio mimicry supergene provide insight into the formation, maintenance and evolution of supergenes. This article is part of the theme issue 'Genomic architecture of supergenes: causes and evolutionary consequences'.

Cover Caption

Butterflies are considered bioindicators of environmental change. We evaluated the combined effects of climate and land‐use interactions on butterfly diversity in tropical rainforest and savanna ecosystems of southwestern China. The results showed that increasing land‐use intensity not only caused a dramatic decrease in butterfly alpha diversity but also significantly simplified butterfly species composition, and finally revealed interactions between human‐modified habitat change and climate change shaped butterfly diversity in both ecosystems. Our study has important implications for biodiversity conservation under the current era of rapid human‐induced habitat loss and climate change (see pages 1109‐1120). The photo is provided by Bai‐Ge Miao, which the butterfly Papilio memnon is visiting the Ixora casei ‐Super king‐ flowers.

Butterfly identification using gray level co-occurrence matrix (glcm) extraction feature and k-nearest neighbor (knn) classification

Gita Persada Butterfly Park is the only breeding of engineered in situ butterflies in Indonesia. It is located in Lampung and has approximately 211 species of breeding butterflies. Each species of Butterflies has a different texture on its wings. The Limited ability of the human eye to distinguishing typical textures on butterfly species is the reason for conducting a research on butterfly identification based on pattern recognition. The dataset consists of 600 images of butterfly’s upper wing from six species: Centhosia penthesilea, Papilio memnon, Papilio nephelus, Pachliopta aristolochiae, Papilio peranthus and Troides helena. The pre-processing stage is conducted using scaling, segmentation and grayscale methods. The GLCM method is used to recognize the characteristics of butterfly images using pixel distance and Angular direction 0o, 45o, 90o and 135o. The features used is angular second moment, contrast, homogeneity and correlation. KNN classification method in this study uses k values1, 3, 5, 7, 9, 11, 13, 15, 17, 19, 21 and 23 based on the Rule of Thumb. The result of this study indicate that Centhosia penthesilea and Papilio nephelus classes can be classified properly compared to the other 4 classes and require a classification time of 2 seconds at each angular orientation. The highest accuracy is 91.1% with a value of in the angle of 90o and error rate8.9%. Classification error occured because the value of the test data features is more dominant with the value of the training image features in different classes than the supposed class. Another reason is because of imperfect test data.

The life cycle of great Mormon butterfly (Papilio memnon memnon) in captivity

Papilio memnon memnon is one of the butterflies groups of the Papilio clan in West Java, Indonesia. This study aims were to determine the morphology, behavior, and life cycle of the P. m memnon butterfly. The research method used is descriptive with survey techniques and literature studies. This study aims to determine the morphology, behavior, and life cycle of the P. m memnon butterfly. The research method used was descriptive with survey techniques and literature studies. Data about morphology, duration of life, and behavior at each development phase during the P. m memnon butterfly life cycle were analyzed descriptively based on direct observation. The results showed the egg phase of P. m. memnon takes about 7-8 days to the phase instar1 of the larvae. The average total time needed in the larval phase is around 23 - 45 days to the pupa phase. The Pre-pupa phase lasts 1-2 days, while the pupa phase lasts 2-3 weeks. P. m. memnon performs the larval phase as many as 5 stages of instar. Food eaten by first phase instar 1 of larvae is the eggshell, which is useful for insect growth and helps eliminate evidence of eggs from parasitic attacks. At each phase of development, the larvae eat the outer skin to obtain additional energy to support growth.

JENIS KUPU-KUPU PAPILIONIDAE DI ZONA PENERIMA TAMAN HUTAN RAYA (TAHURA) RAJO LELO BENGKULU

This study aims to determine the type of butterfly Papilionidae in the Receiver Zone of the Great Forest Park (GFP) Raja Lelo Bengkulu. The study was conducted in June- August 2017. Determination of the location of the study was conducted using purposive sampling and butterfly collection at each location using the roaming method. Butterfly collection is carried out at 08.00-10.00 WIB and continued in the afternoon at 15.00- 17.00 WIB, on sunny weather using insect nets. The butterflies obtained were taken to the Biology Laboratory of the Basic Science building, then stretched on a stretching board and dried in an oven at 40 0 C. Furthermore, the butterfly sample is identified by the butterfly identification book and grouped by Family, Genus and Species. The data obtained were analyzed descriptively. The results showed 7 species of Papilionidae butterflies, 74 individuals belonging to 4 genera. The types of butterflies obtained were Graphium agamemnon, Grapihum sarpedon, Pachliopta aristolochiae, Papilio polytes, Papilio memnon, Papilio palinurus, Troides helena.

Identifikasi Kupu-Kupu Menggunakan Ekstraksi Fitur Deteksi Tepi (Edge Detection) dan Klasifikasi K-Nearest Neighbor (KNN)

Lampung has the only breeding of in situ butterflies engineered in Indonesia namely Gita Persada Butterfly Park, which has approximately 211 butterfly species. Butterflies can be classified according to patterns found on the wings of a butterfly. The weakness of the human eye in distinguishing patterns on butterflies is a foundation in building butterfly identification based on pattern recognition. This study uses 6 species of butterflies: Papilio memnon, Troides helena, Papilio nephelus, Cethosia penthesilea, Papilio peranthus, and Pachliopta aristolochiae. The butterfly dataset used is 600 images. The butterfly image used is in the form of the upper wing side. The pre-processing stage uses the method of scaling, segmentation, and grayscale. The feature extraction stage uses the canny edge detection method by applying smoothing, edge strength, edge direction, non maximum suppression, and hyterisis thresholding. The classification phase uses the K-Nearest Neighbor (KNN) method with values k = 1, 3, 5, 7, 9, 11, 13, 15, 17, 19, 21 and 23 obtained under the Rule of Thumb. The identification of butterfly require a classification time of 8 seconds. The highest accuracy is obtained from testing with a value of k = 5 by 80%.

Feeding Preference of Papilio memnon Linnaeus, 1758 (Lepidoptera) Larvae on Host Plants Citrus aurantifolia and Citrus hystrix (Rutaceae)

This research aimed to investigate the food preference of the Papilio memnon L. larvae on two host plants. This study was conducted by two choice food preference test combination of leaves Citrus aurantifolia Chrism. and Citrus hystrix DC. The result of this study showed that the third instar of P. memnon L. larva significant preferential feeding (P<0.05), while the fourth and fifth instar shown that no significant feeding preference beetwen host plants. Nitrogen and water content of leaves were higher on C. aurantifolia (0.212 % and 70.368%) than C. hystrix (0.04% dan 64.027%). While carbohydrate content was higher on C. hystrix (22.167%) than C.aurantifolia (31.097%).

Larval Growth of Great Mormon Butterfly (Papilio memnon memnon) Fed with Citrus aurantifolia Leaves

Papilio memnon memnon is one of butterflies that has a specific host plant to lay eggs and for its larval growth, one of the host plants is a Citrus. Citrus leaf plays an important role for the growth of butterflies during the larval phase. The aim of this research is to know nutritional composition of C. aurantifolia leaves, the growth of Papilio memnon memnon larvae which is fed with lime leaf (Citrus aurantiifolia) on Laboratory scale and other factors that influences its growth. The research method used was experiment, design of Completely Random Design with larvae feed as independent variable and larvae growth as dependent variable. The data were analyzed descriptively including body length, larvae body weight, and head capsule width followed by ANOVA and t-test. The results showed that the feeding of C. aurantiifolia leaves had no significant effect on body weight or body length of P. m memnon in all larval phases, from instar 1 to instar 5, but significantly affect the width of larval capsule cap at instar 4 and instar to 5. In the development phase, P. m. memnon larvae found symptoms of Nuclear Polyhedrosis Virus (NPV) disease that causes the death of larvae.

Parallel evolution of Batesian mimicry supergene in two Papilio butterflies, P. polytes and P. memnon.

Batesian mimicry protects animals from predators when mimics resemble distasteful models. The female-limited Batesian mimicry in Papilio butterflies is controlled by a supergene locus switching mimetic and nonmimetic forms. In Papilio polytes, recent studies revealed that a highly diversified region (HDR) containing doublesex (dsx-HDR) constitutes the supergene with dimorphic alleles and is likely maintained by a chromosomal inversion. In the closely related Papilio memnon, which exhibits a similar mimicry polymorphism, we performed whole-genome sequence analyses in 11 butterflies, which revealed a nearly identical dsx-HDR containing three genes (dsx, Nach-like, and UXT) with dimorphic sequences strictly associated with the mimetic/nonmimetic phenotypes. In addition, expression of these genes, except that of Nach-like in female hind wings, showed differences correlated with phenotype. The dimorphic dsx-HDR in P. memnon is maintained without a chromosomal inversion, suggesting that a separate mechanism causes and maintains allelic divergence in these genes. More abundant accumulation of transposable elements and repetitive sequences in the dsx-HDR than in other genomic regions may contribute to the suppression of chromosomal recombination. Gene trees for Dsx, Nach-like, and UXT indicated that mimetic alleles evolved independently in the two Papilio species. These results suggest that the genomic region involving the above three genes has repeatedly diverged so that two allelic sequences of this region function as developmental switches for mimicry polymorphism in the two Papilio species. The supergene structures revealed here suggest that independent evolutionary processes with different genetic mechanisms have led to parallel evolution of similar female-limited polymorphisms underlying Batesian mimicry in Papilio butterflies.

Diversity of Species and Conservation Priority of Butterfly at Suranadi Natural Park of West Lombok, Indonesia

Butterflies play an important role in the ecosystem of Suranadi Natural Park in West Lombok. Butterflies help preserving the existence and diversity of flora by facilitating the process of pollination so it is crucial for flowering plants. The present study aimed at analyzing the diversity and determining the priority of butterfly conservation at Suranadi Natural Park of West Lombok. This exploration is a descriptive study. Data were observed in four-time repetition in the morning and in the afternoon for two months. The sweeping net technique following observation path was employed in this research (line left, line right, line central and line waterway). The data analysis used the Shannon-Wiener diversity index and priority for conservation determined by Ministry of Forestry Regulation Number: P.57 / Menhut-II / 2008. Fourty (40) species of butterflies belonging to 5 families identified. The diversity index (H ') of butterflies ranges from 2.63 to 3.43 (medium-high). The two species of the Papilionidae family found at TWA Suranadi, namely, Troides helena and Papilio memnon were the priority of conservation. This research would be helpful to consider conservation strategy and ecotourism.

Do juvenile developmental and adult body characteristics differ among genotypes at the doublesex locus that controls female-limited Batesian mimicry polymorphism in Papilio memnon?: A test for the “cost of mimicry” hypothesis

Female-limited Batesian mimicry may have evolved because of stronger predation pressure on females than on males, but some physiological costs of mimicry may also hinder the evolution of mimicry in males. In Papilio memnon, which possesses a female-limited Batesian mimicry polymorphism, two alleles at the doublesex (dsx) locus strictly control female phenotypes. To examine whether there are physiological costs associated with mimetic genotypes in the juvenile stage, we compered mortality, juvenile growth and development, and the resultant adult characteristics among three dsx genotypes (HH, Hh, hh) at a constant temperature (25 °C) and two differing day lengths (LD 14:10 and LD 12:12; the latter might induce pupal diapause) by crossing individuals heterozygous (Hh) for the dsx allele. All pupae emerged directly without diapause irrespective of day length. The genotype frequencies of the emerged individuals were consistent with the expected 1:2:1 ratio of HH:Hh:hh. The sex ratio was significantly male-biased in one of two families, but not in the other. We found no effect of genotype on any developmental or adult characteristic, although there were sex differences in most traits. The larval development time was longer and growth rate higher in females than in males; pupal weight, forewing length, and total dry mass of the thorax and abdomen were greater in females, whereas the thoracic mass/abdominal mass ratio was greater in males. We also found that the growth rate was higher and pupal period longer with a short day than with a long day. Overall, we found no evidence for physiological costs associated with the mimetic genotypes. However, it is too early to conclude that no physiological cost of mimicry affects the evolution and maintenance of this female-limited Batesian mimicry polymorphism because we have not studied the adults of different genotypes.

Temporal dynamics of the mimetic allele frequency at the doublesex locus, which controls polymorphic Batesian mimicry in Papilio memnon butterflies

Tracking allele frequencies is essential for understanding how polymorphisms of adaptive traits are maintained. In Papilio memnon butterflies, which exhibit a female-limited Batesian mimicry polymorphism (wing-pattern polymorphism), two alleles at the doublesex (dsx) locus correspond to mimetic and non-mimetic forms in females; males carry both dsx alleles but display only the non-mimetic form. This polymorphism is thought to be maintained by a negative frequency-dependent selection. By tracking dsx allele frequencies in both sexes at a Taiwanese site over four years, we found that the mimetic allele persists at intermediate frequencies even when the unpalatable model papilionid butterflies (Pachliopta and Atrophaneura species) were very rare or absent. The rates of male mate choice did not differ between the two female forms; neither did insemination number nor age composition, suggesting equivalent reproductive performance of the two forms over time. Our results characterised the temporal dynamics of the mimetic allele frequency in the field for the first time and give insights into underlying processes involved in the persistence of the female-limited Batesian mimicry polymorphism.