Diapause pupal color diphenism induced by temperature and humidity conditions in Byasa alcinous (Lepidoptera: Papilionidae)

We investigated whether diapause pupae of Byasa alcinous exhibit pupal color diphenism (or polyphenism) similar to the diapause pupal color polyphenism shown by Papilio xuthus. All diapause pupae of B. alcinous observed in the field during winter showed pupal coloration of a dark-brown type. When larvae were reared and allowed to reach pupation under short-day conditions at 18 °C under a 60 ± 5% relative humidity, diapause pupae exhibited pupal color types of brown (33%), light-brown (25%), yellowish-brown (21%), diapause light-yellow (14%) and diapause yellow (7%). When mature larvae reared at 18 °C were transferred and allowed to reach pupation at 10 °C and 25 °C under a 60 ± 5% relative humidity after a gut purge, the developmental ratio of brown and light-brown, yellowish-brown, and diapause light-yellow and diapause yellow types was 91.2, 8.8 and 0.0% at 10 °C, and 12.2, 48.8 and 39.0% at 25 °C, respectively. On the other hand, when mature larvae reared at 18 °C were transferred and allowed to reach pupation at 10 °C, 18 °C and 25 °C under an over 90% relative humidity after a gut purge, the developmental ratio of brown and light-brown, yellowish-brown, and diapause light-yellow and diapause yellow types was 79.8, 16.9 and 3.3% at 10 °C, 14.5, 26.9 and 58.6% at 18 °C, and 8.3, 21.2 and 70.5% at 25 °C, respectively. These results indicate that diapause pupae of brown types are induced by lower temperature and humidity conditions, whereas yellow types are induced by higher temperature and humidity conditions. The findings of this study show that diapause pupae of B. alcinous exhibit pupal color diphenism comprising brown and diapause yellow types, and suggest that temperature and humidity experienced after a gut purge are the main factors that affect the diapause pupal coloration of B. alcinous as environmental cues.     

Population dynamics ofLuehdorfia japonica Leech (Lepidoptera: Papilionidae)

1. An adult population of a papilionid butterfly, Luehdorfia japonicaLeech , was studied by marking, release and recapture procedures in a hilly region in the suburbs of Kanazawa City, Japan.
2. Age of butterflies was estimated from the wing wear conditions, rated as winage categories 0 to 6.
3. Jolly (1965) andSeber's (1973) method was applied to the marking-recapture results for estimating the population parameters (sampling ratio, population size and survival rate).
4. Sampling ratio of males was consistently higher (around 50%) than that of females.
5. Newly emerged females were especially inactive, so that few of them were captured. From day 6.5 to day 10.0 they began to oviposit and became more active and more catchable.
6. An approximate sex ratio of 1∶1 was confirmed from the specimens collected in the field and by rearing experiments.
7. Daily survival rate was about 0.75–0.80 and mean longevity was about 4 days for both sexes. The maximum longevity observed was 17 days, for males and 21 days for females.
8. Dispersal by both sexes of the butterfly was more than 1 km.

     

Environmental factors affecting black/white coloration of the silken girdle in the swallowtail butterfly, Atrophaneura alcinous (Lepidoptera: Papilionidae)

The silken girdles of pupae of the swallowtail butterfly Atrophaneura alcinous show black and white color diphenism. Field observations revealed that all pupae observed on non-food plants and the leaves and stems of the larval food plant Aristolochia debilis were classified as a silken girdle of a black type, while a large portion of pupae pupating on the twigs and trunks of cherry trees in close proximity to A. debilis were classified as a silken girdle of a black type. Additionally, all pupae observed on the surfaces of artificial objects in areas where there are no surrounding plants or trees were classified as a silken girdle of a white type. We demonstrated the effect of day length and the texture, light, plant odor and humidity of pupation sites on the coloration of the silken girdle in A. alcinous. Regardless of long-day or short-day day length conditions, light conditions of constant light or dark, or the presence of a plant odor of A. debilis as environmental cues, all larvae placed at over 80% relative humidity (R.H.) developed into pupae with a silken girdle of a black type. However, all larvae developed into pupae with a silken girdle of a white type when R.H. was below 75%. Furthermore, when pupae with a silken girdle of a white type were transferred to conditions of 90% R.H. within 24 hr of pupation, the white color of the silken girdle changed into a black type within 24 hr of the transfer. The present data suggest that the induction of a black coloration of the silken girdle in A. alcinous requires a R.H. of approximately 80% or more as an environmental factor.     

Population dynamics ofLuehdorfia japonica Leech (Lepidoptera: Papilionidae)

Mortality processes from egg to final instar larval stage are examined in the papilionid butterfly Luehdorfia japonica for two generations in a natural populations. Special attention is given to the effects of initial egg cluster size on the survival rate by the 3rd instar. Mean egg cluster sizes for the two generations were 11.3 and 10.7. The hatching rate was not affected by the egg cluster size. Up to 3rd instar, larvae in a group were more likely to die en masse rather than to die individually, whereas most larvae in later instars died independently. The egg clusters of the average (and the most frequent) size class (11±1) had the second highest survival rate until the 3rd instar. A slightly higher survival rate occurred in clusters a little larger (14±1). The average size clusters were least likely to be exterminated and about 70% of them produced at least one individual surviving to the 3rd instar. Factors affecting the observed mortality patterns were discussed.     

Search behavior of pheromone-stimulated potato tuber moth males (Lepidoptera: Gelechiidae)

The processes of female searching by male potato tuber moths,Phthorimaea operculella, were analyzed. The behavioral components to copulation were antennal cleaning, quiescence, walking, wing fanning, contact with female, hair brush display, copulation attempt, and copulation. Males did not always succeed in mating on their first attempt. Searching behavior of males changes to “area-restricted searching” after contact with a female. Males could, therefore, find females efficiently and copulate.     

Comparative processing of allelochemicals in the Papilionidae (Lepidoptera)

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

A partitioned likelihood analysis of swallowtail butterfly phylogeny (Lepidoptera:Papilionidae)

Although it is widely agreed that data from multiple sources are necessary to confidently resolve phylogenetic relationships, procedures for accommodating and incorporating heterogeneity in such data remain underdeveloped. We explored the use of partitioned, model-based analyses of heterogeneous molecular data in the context of a phylogenetic study of swallowtail butterflies (Lepidoptera: Papilionidae). Despite substantial basic and applied study, phylogenetic relationships among the major lineages of this prominent group remain contentious. We sequenced 3.3 kb of mitochondrial and nuclear DNA (2.3 kb of cytochrome oxidase I and II and 1.0 kb of elongation factor-1 alpha, respectively) from 22 swallowtails, including representatives of Baroniinae, Parnassiinae, and Papilioninae, and from several moth and butterfly outgroups. Using parsimony, we encountered considerable difficulty in resolving the deepest splits among these taxa. We therefore chose two outgroups with undisputed relationships to each other and to Papilionidae and undertook detailed likelihood analyses of alternative topologies. Following from previous studies that have demonstrated substantial heterogeneity in the evolutionary dynamics among process partitions of these genes, we estimated evolutionary parameters separately for gene-based and codon-based partitions. These values were then used as the basis for examining the likelihoods of possible resolutions and rootings under several partitioned and unpartitioned likelihood models. Partitioned models gave markedly better fits to the data than did unpartitioned models and supported different topologies. However, the most likely topology varied from model to model. The most likely ingroup topology under the best-fitting, six-partition GTR + gamma model favors a paraphyletic Parnassiinae. However, when examining the likelihoods of alternative rootings of this tree relative to rootings of the classical hypothesis, two rootings of the latter emerge as most likely. Of these two, the most likely rooting is within the Papilioninae, although a rooting between Baronia and the remaining Papilionidae is only nonsignificantly less likely.     

Male-like females, mimicry and transvestism in butterflies (Lepidoptera: Papilionidae)

Abstract. When a butterfly species has a polymorphic female, with one of the forms closely resembling the male, it is customary to suppose that this form is ancestral, and that the ‘odd’ forms have arisen later. R. I. Vane-Wright, on the other hand, has suggested that in some species the male-like form may be a ‘transvestite’ female, the ancestral form of the female having been strikingly unlike the male. As later-derived forms are usually, but not always, genetically dominant to ancestral forms, we can make some choice between these hypotheses by discovering the dominance relations of the male-like and the ‘odd’ forms of the female. In the mimetic Papilio aegeus the male-like form is shown to be recessive to the ‘odd’ (mimetic) form, as has essentially been the case in all other butterflies so far investigated. Papilio phorcas is now shown to be the exception: the ‘odd’ (non-mimetic) form is recessive to the male-like form. We conclude that usually the male-like form is ancestral, but that P.phorcas may be an authentic example of ‘transvestism’, or the ‘transfer’ of male epigamic colour to the female of the species. The yellow, male-like pattern of the mimetic Papilio dardanus may be dominant or recessive to the mimetic forms according to the genetic background: largely recessive in Madagascar, and southern and western Africa, dominant to most forms in Ethiopia, and probably dominant to one mimetic form but recessive to the others in Kenya. All female dardanus patterns, both mimetic and yellow, are strongly dominant to both female phorcas patterns in P. dardanus × P. phorcas hybrids (P. ‘nandina’). The simplest explanation of this situation is that the male-like pattern of dardanus is ancestral, and that dominance has become locally reversed in Ethiopia. The dominance relations, and the sex- or autosomal-linkage of two forms can be determined without pedigree-breeding, simply by observing a few offspring each from a large number of wild-caught females.     

Speciation problems in the Papilio machaon group of butterflies (Lepidoptera: Papilionidae)

Abstract. The taxonomic and genetic relationships between P.machaon rathjensi Warnecke from the west side of southern Arabia and P.machaon muetingi Seyer from the east are discussed, together with suggestions of their relationship to P.machaon saharae Oberthiir from the northern Hejaz, Sinai and desert North Africa. It is concluded that though P.m.rathjensi and P.m.saharae are subspecifically distinct, they are very closely related to each other, and both should be regarded as specifically distinct from the rest of the machaon complex.
The remainder of the machaon complex in its turn presents a somewhat similar problem. Thus P.machaon gorganus Fruhstorfer and P.hospiton Géné are normally regarded as good species, but Fl caterpillars have been found in the wild and the cross can be readily produced in captivity with no upset in the sex ratio. Furthermore, recently we have obtained sparse F2 offspring between the two species.
We therefore suggest that the entire P.machaon complex is in a labile state as regards speciation throughout its range and the relationships between the various forms are not easily expressed through the traditional species and subspecies concepts. Our tentative conclusions are that the status of P.machaon and P.hospiton as distinct species probably remains valid. The saharaelrathjensi complex might be considered distinct from P.machaon using the biological species concept, while their genetics indicate a more conservative approach. However, more important than the exact status of the taxa in question is the fact that the P.machaon complex illustrates evoluton in action, the end result of which cannot be predicted.     

Soldier bug predation on swallowtail caterpillars (Lepidoptera: Papilionidae): Circumvention of defensive chemistry

All larvae in the family Papilionidae possess osmeteria, eversible glands that release defensive chemicals upon disturbance. These chemicals have been shown to be repellent to ants, mantids, and other predaceous arthropods with chewing mouthparts. In this study, we demonstrate that the pentatomid Podisus maculiventris,an important predator of the black swallowtail Papilio polyxenes,is capable of piercing the body wall of the caterpillar and consuming body fluids without eliciting osmeterial eversion. Tenebrio molitorlarvae coated with osmeterial secretions of Papilio polyxeneswere universally rejected by Podisus maculiventris,suggesting that the predator is in fact repelled by the swallowtail defensive chemistry. By circumventing the defensive behavior of the caterpillar, the pentatomid is able to utilize an otherwise chemically unsuitable prey species.     

Effects of plant odor on oviposition by the black swallowtail butterfly,Papilio polyxenes (Lepidoptera: Papilionidae)

Black swallowtail females laid more eggs on plant models treated with contact stimulants and volatiles from carrot leaves than on models treated only with contact stimulants. The volatiles enhanced landing rates and females alighted more frequently on artificial leaves treated with host volatiles than on adjacent control leaves. Volatiles from cabbage, a nonhost, inhibited landing rates on artificial leaves treated with carrot contact stimulants. Examination of antennae revealed two major types of sensilla, believed to be olfactory in function. Electroantennogram preparations responded more strongly to carrot volatiles than to cabbage volatiles and several shared responses at particular retention times to carrot volatile components eluting from a gas chromatograph. Our results are consistent with a long-standing hypothesis that behavioral responses to essential oil components characteristic of the larval food plants have facilitated host shifts in the genus Papilio.     

The Effect of Metals on the Mortality of Parnassius Apollo Larvae (Lepidoptera: Papilionidae)

The population sizes and range of Parnassius apollo started to decline in Finland in the 1930s and resulted in complete extinction in many parts of its former range 30 years later. It has been suggested that metal pollution is one potentially very important reason for the decline. Recently, a recovery of P. apollo and widening of its range has started in the southwestern Finland, simultaneously with a decline in metal fallout from air. Metal analyses were performed with larval and host plant samples collected from an area where P. apollo is recolonising its former range (the archipelago between the island of Kemiö and the Hankoniemi peninsula). In the host plants (Sedum telephium), the level of Fe was significantly lower and the levels of Cd and Zn suggestively lower in the current range (n = 35) of P. apollo than in the former range (n = 12). Analyses on larvae of P. apollo (n = 29) showed lower levels of Cd, Fe and Mn than in the host plant leaves. The excretion of these metals, as well as Zn, via the faeces was effective. The level of the Cd-antagonist Cu was considerably higher in caterpillars than in the food. When caterpillars were fed with metal-polluted food from Helsinki city (4.35ppm Cd/dwt; n = 5) they died, whereas control individuals fed with food from the archipelago (2.49ppm Cd/dwt; n = 5) survived. These results indicate that metals may play an important role in the large-scale decline and partial recovery of P. apollo in the western Europe.     

Temperature-dependent development of overwintering Sericinus montela Gray (Lepidoptera: Papilionidae) pupae and its validation

The developmental time and survival of overwintering Sericinus montela Gray pupae were studied at four constant temperatures (15.0, 20.0, 25.0, and 30.0 °C), 40 ± 10% relative humidity, and 10:14 h light:dark cycle. The developmental time of both sexes decreased with increasing temperature between 15.0 °C (70.18 days for females and 55.28 days for males) and 30.0 °C (19.60 days for females and 13.78 days for males). The development periods of females were longer than those of males at each constant temperature. The relationship between the developmental rate and temperature was fitted by a linear model and a nonlinear developmental rate model (Lactin 1). The mortality of overwintered S. montela pupae was lowest at 25.0 °C (16.7%) and highest at 15.0 °C (36.7%). The lower developmental thresholds were 12.38 and 12.16 °C for females and males, respectively. The distribution of development completion for females and males was described by the two-parameter Weibull distribution equation (r² = 0.87 for females and r² = 0.94 for males). The date for the cumulative 50% adult emergence was within one or two days of that predicted using the Lactin 1 model. The temperature-dependent developmental model for S. montela could be applied to predict the timing of spring emergence in different geographical locations and will be helpful in developing a full-cycle phenology model for S. montela.     

Papilio aegeus Donovan (Lepidoptera: Papilionidae) host plant range evaluated experimentally on ancient angiosperms

Abstract  Chemical similarities among ancient Angiosperms presumably played a role in the ecological and evolutionary diversification of the swallowtail butterflies (Papilionidae). The abilities of neonate larvae of the Citrus swallowtail, Papilio (= Princeps ) aegeus (from Queensland, Australia), to eat, survive and grow on leaves (a choice of young and old) of 34 plant species from families of ancient Angiosperms; 8 Rutaceae, 3 Magnoliaceae, 13 Lauraceae, 3 Monimiaceae, 1 Aristolochiaceae, 2 Apiaceae, 1 Sapotaceae, 1 Winteraceae and 2 Annonaceae were tested. It was apparent that there is genetic variation in populations of Rutaceae-specialised Australian P. aegeus for acceptance, consumption and larval growth, reflecting differential suitability of some native Australian Lauraceae species as food plants (as well as certain Winteraceae, Monimiaceae and non-Australian Magnoliaceae, Lauraceae and Annonaceae). No consumption or survival of P. aegeus was seen on Aristolochia elegans (Aristolochiaceae) or Pouteria australis (Sapotaceae) despite literature records alluding to this possibility. The Rutaceae specialist P. aegeus appears to have the fundamental detoxification capabilities for processing many existing species of the basal Angiosperm families, without having direct ancestors that historically had fed on them.     

Towards resolving the identities of the Graphium butterflies (Lepidoptera: Papilionidae) of Peninsular Malaysia

Graphium butterflies are famous in Peninsular Malaysia for their colourful wings, yet their taxonomy remains unresolved. The popular guides to Malaysian butterflies, place the species in one, two or three genera and identification to species using obscure morphological characters can be difficult, especially for the closely related species, G. bathycles bathycloides and G. chironides malayanum. We sequenced the COI mtDNA barcode for Graphium specimens in the Museum of Zoology, University of Malaya to test the utility of DNA barcoding for the identification of Graphium species. Additionally, we sequenced 28S rRNA to examine, in conjunction with COI, the phylogenetic relationships of these species and investigate the validity of Pathysa and Paranticopsis as distinct genera. We found that all species of Graphium possessed a distinctive cluster of DNA barcodes with the exception of specimens originally identified as G. bathycles bathycloides and G. chironides malayanum which shared DNA barcodes. On further examination we found that the morphological determinations were ambiguous as the specimens overlapped for diagnostic characters reported for each species. The COI and 28S rRNA phylogenetic trees showed a similar topology with Paranticopsis species forming a clade nested within a larger clade also comprising Pathysa species. Based on this topology, in order for Pathysa to be a valid genus, at least three other clades within Graphium s.l. would also have to be raised as genera.     

Wing colors of Chrysozephyrus butterflies (Lepidoptera; Lycaenidae): ultraviolet reflection by males

Wing colors of the four species of Chrysozephyrus butterflies were analyzed by a spectrophotometer. As the dorsal wing surface of males showed a strong reflectance when the specimen was tilted, measurements were made by the tilting method. The dorsal wing surface of males which appears green to the human eye reflected UV (315-350 nm) as well as green light (530-550 nm). The reflectance rate of UV to visible green light varied among species with a higher rate for C. hisamatsusanus and C. ataxus, and a lower rate for C. smaragdinus and C. brillantinus. The peak wavelength and the peak height did not shift when the specimen was exposed to direct sunlight at least for 16 hr. Artificial removal of scales by scratching the wing surface decreased reflectance. Blue marks on the forewings of C. brillantinus, C. hisamatsusanus and C. ataxus females reflected UV to visible light of short wavelength, and orange marks on the dorsal surface of the forewing and the ventral surface of the hindwing of C. samaragdinus females showed a higher reflectance at longer wavelengths.     

The complete mitochondrial genome of the endangered Apollo butterfly,Parnassius apollo (Lepidoptera: Papilionidae) and its comparison to other Papilionidae species

The Apollo butterfly, Parnassius apollo is a representative species of the butterfly subfamily Parnassiinae. This charming species is one of the most endangered butterfly species in the world. In this study, we sequenced its complete mitochondrial genome (mitogenome), with the aim of accumulating genetic information for further studies of population genetics and mitogenome evolution in the Papilionidae. The 15,404-bp long mitogenome harbors a typical set of 37 genes and is the largest butterfly mitogenome determined, except for Papilio maraho (16,094 bp). Like many other sequenced lepidopteran species, one tRNA^Trp-like and one tRNA^Leu(UUR)-like sequences were detected in the AT-rich region. A total of 164 bp of non-coding sequences are dispersed in 14 regions throughout the genome. The longest intergenic spacer (68 bp) is located between tRNA^Ser(AGN) and tRNA^Glu, and is the largest spacer at this location among Papilionidae species. This spacer may have resulted from an 8-fold repetition of a TTTCTTCT motif or a 4-fold repetition of a CTTTATTT motif.     

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

Abstract.

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