Geographic variation in photoperiodic diapause induction and diapause intensity in Sericinus montelus (Lepidoptera: Papilionidae)

Abstract Due to the risk of extinction and ornamental value of the swallowtail butterfly, Sericinus montelus Gray (Lepidoptera: Papilionidae) in China, knowledge about local adaptations is important for the conservation and economical utilization of the species. In the present study, photoperiodic diapause induction and diapause intensity of S. montelus populations from Jiamusi (46°37′N), Beijing (40°15′N), Zibo (36°48′N), Fangxian (32°36′N), Wuhan (30°33′N) and Huaihua (27°33′N) were characterized at 25°C. Logistic regression analysis revealed a significant population × hours of light interaction, confirming that photoperiodic responses varied among populations. The critical photoperiod was positively correlated with latitude and increased toward the north at a rate of about 1 h for each 6.67 degrees of latitude. Survival analyses indicated that survival time of diapausing pupae before adult eclosion differed significantly among populations at 25°C and 16 : 8 L : D h. The mean duration of pupal diapause was also positively correlated with latitude. Our study reveals geographic variation in the critical photoperiod for diapause induction and in diapause intensity of S. montelus. These results provide useful information for our general understanding about seasonal adaptation in insects and may also be used to predict how geographic populations respond to climate warming.     

Effect of soil moisture on overwintering pupae in Spodoptera exigua (Lepidoptera: Noctuidae)

Spodoptera exigua Hübner (Lepidoptera: Noctuidae), can overwinter as pupae at depths of 0- to 5-cm beneath the ground. In order to understand the effect of soil moisture on the overwintering success of this species, we tested the supercooling point and survival of pupae, and the growth rate of prepupae that were exposed to different temperature and soil moisture in the laboratory. Results showed that supercooling points, body water contents, and survivals after the different pre-treatments were not significantly affected by the soil moisture. The developmental progress of prepupae, survival of prepupae and pupae were negatively correlated with soil moisture as well as the delay of exposure time, which supports the hypothesis that soil moisture acts as a developmental modulator. We presumed that the delay of pupation and lower prepupae survival under higher soil moisture was due to lack of a complete pupal chamber for protection at low temperatures. Low pupal survival was likely attributed to lack of oxygen in the soil, especially under the condition of higher soil moisture. We suggest that using strategies of irrigation and soil tillage during winter may decrease the overwintering population of S. exigua from the perspective of integrated pest management.     

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.

     

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.     

Pair-clinging behavior by males ofAtrophaneura alcinous (Lepidoptera: Papilionidae)

Females ofAtrophaneura alcinous usually mate soon after eclosion. Their ostium bursae becomes plugged with male secretion which reduces chances of remating. Males frequently cling to a copulating pair and wait for completion of copulation. This was observed in 66% of 198 copulating pairs, with a maximum of 5 males clinging at one time during the course of a copulation. Males clinging for longer periods were more successful in copulation with the freshly mated female than those clinging for shorter periods. Despite the plugging effect, females may mate more than once. Clinging males were responsible for 61% of re-copulations and 53% of re-inseminations. Clinging behavior may be regarded as an effective male mating strategy to exploit freshly mated females, and an alternative to finding virgin females.     

Mobility and oviposition site-selection in Zerynthia cassandra (Lepidoptera,Papilionidae): implications for its conservation

The adults’ mobility and oviposition preferences of Zerynthia cassandra have been studied for the first time, with the aim of integrating auto-ecological information into recommendations for the habitat’s management of this species. Results of our mark-release-recapture study have highlighted that Z. cassandra is a strictly sedentary species, since detected movements only occurred over very short distances (≤200 m) and mainly within the species’ reproductive habitat (i.e. around Aristolochia rotunda stands), with males moving further than females. Our study shows that the main oviposition habitat of Z. cassandra is found where A. rotunda plants are growing in large stands; sites where plants growing in half to full sun and mostly oriented to the south are preferred. The distance of deposited eggs from the plants’ roots was narrowly correlated with the plants’ length. Eggs were deposited singly, mainly on the underside of leaflets. Management strategies necessary for improving the most important habitat features for the conservation of this species are suggested.     

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.     

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.     

Physiology of diapause and cold hardiness in the overwintering pupae of the fall webworm Hyphantria cunea (Lepidoptera: Arctiidae) in Japan

The fall webworm Hyphantria cunea Drury, which was accidentally introduced to Japan in 1945, overwinters on the ground in pupal diapause. Diapause termination, as indicated by the respiration rate and the period required for adult emergence, began in March and ended in April. Cold hardiness (the ability to survive exposure to -15 degrees C) decreased linearly with diapause development from November to the following April under field conditions. Cold hardiness of diapause pupae (DP) decreased as the acclimation temperature decreased from 15 to -10 degrees C, whereas cold hardiness of non-diapause pupae (NDP) remained high as the acclimation temperature decreased from 5 to -5 degrees C. However, H. cunea in Japan can survive exposure to -5 degrees C for two weeks, whether it is in a diapause or non-diapause state. Trehalose was the main sugar detected in the body, but its level was less than 0.8%. Trehalose levels increased in field-collected pupae from January to March. DP accumulated less trehalose than NDP, as the acclimation temperature was decreased from 5 to -5 degrees C. The alanine content in field-collected pupae increased from November to February. Both diapause and low temperature caused an accumulation of alanine. These results suggest that under field conditions, overwintering pupae of H. cunea in Japan do not accumulate high levels of sugars and polyols and do not develop a high level of cold hardiness. Furthermore, DP do not accumulate high levels of sugars and polyols and their ability to survive exposure to -15 degrees C is not greater than that of NDP. The physiological and biochemical bases of diapause in H. cunea from Japan are discussed.     

Hormonal control of the orange coloration of diapause pupae in the swallowtail butterfly, Papilio xuthus L. (Lepidoptera: Papilionidae)

Diapause pupae of Papilio xuthus show color polymorphism, represented by diapause-green, orange, and brownish-orange types that are each associated with specific pupation sites. We investigated the role of the site of pupation on the induction of the development of orange types (or brownish-orange types), and the endocrine mechanism underlying the control of color polymorphism in short-day pupae. All short-day larvae of the wandering stage developed into orange or brownish-orange type pupae when they were placed in rough-surfaced containers after gut-purge. Utilizing a pharate pupal ligation between the thorax and abdomen, the endocrine mechanism underlying the control of color polymorphism was shown to involve a head-thorax factor (Orange-Pupa-Inducing Factor: OPIF) that induced orange types in short-day pupae. OPIF was bioassayed using the ligated abdomens of short-day pharate pupae. OPIF was extractable with 2% NaCl solution from 5th-instar larval ganglia complexes following the mesothoracic complex (TG(2,3)-AG(1-7)), but it could not be extracted with either acetone or 80% ethanol solution. OPIF may not exist in the brains of day-0 pupae or in brain-subesophageal ganglion and prothoracic ganglion complexes of 5th-instar larvae. The short-day pharate pupae responded to OPIF in a dose-dependent manner.     

Cold hardiness of Helicoverpa zea (Lepidoptera: Noctuidae) pupae

An insect's cold hardiness affects its potential to overwinter and outbreak in different geographic regions. In this study, we characterized the response of Helicoverpa zea (Boddie) pupae to low temperatures by using controlled laboratory measurements of supercooling point (SCP), lower lethal temperature (LT(50)), and lower lethal time (LLTime). The impact of diapause, acclimation, and sex on the cold hardiness of the pupae also were evaluated. Sex did not significantly affect the SCP, LT(50), or LLTime. However, the mean SCP of diapausing pupae (-19.3°C) was significantly lower than nondiapausing pupae (-16.4°C). Acclimation of nondiapausing pupae to constant temperatures from 10 to 20°C before supercooling also produced a significantly lower SCP than nondiapausing pupae held at 25°C. The LT(50)s of nondiapausing and diapausing were not significantly different, but confirmed that H. zea pupae are chill-intolerant because these lethal temperatures are warmer than the corresponding mean SCPs. Diapausing pupae survived longer than nondiapausing pupae at the same, constant, cold temperatures, a finding consistent with the SCP results. Both of these results suggest enhanced cold hardiness in diapausing pupae. When laboratory results were compared with field temperatures and observed distributions of H. zea in the contiguous United States, the laboratory results corroborated what is currently perceived to be the northern overwintering limit of H. zea; approximately the 40(th) parallel. Moreover, our research showed that areas north of this limit are lethal to overwintering pupae not because of low temperature extremes, but rather the length of time spent at near-zero temperatures.     

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.     

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.     

Temperature-dependent demography of Chilades pandava peripatria (Lepidoptera: Lycaenidae)

Chilades pandava peripatria Hsu and its host plant Cycas taitungensis Shen, Hill, Tsou & Chen are both endemic species to Taiwan. Ch. pandava peripatria has a specific association with buds and soft leaves of cycad plants. The introduced species, Cy. revoluta, have prolonged budding periods and extensive auxiliary buds that extensively contribute to the outbreak of Ch. pandava peripatria. An in-depth knowledge of the development, survival, and fecundity of Ch. pandava peripatria under different environmental conditions is necessary to understand the population growth of Ch. pandava peripatria. The demography of Ch. pandava peripatria was studied based on the age-stage, two-sex life table at 20, 23, 25, 28, and 31 degrees C, 70% RH, and a photoperiod of 16:8 (L:D) h under laboratory conditions. Ch. pandava peripatria completed its development under tested temperatures but did not produce offsprings at 23 degrees C. Because of the high egg mortality at 20 degrees C, the data at this given temperature were excluded from this study. The intrinsic rate of increase (r) under these tested temperatures was 0.1846, 0.2919, and 0.1412 d(-1), respectively. The net reproductive rate (H(o)) was 165.47, 262.32, and 56.68 offsprings per individual and the mean generation time (T) was 27.72, 19.10, and 28.67 d, respectively. Our results indicated that Ch. pandava peripatria is highly adaptable to environments where temperature ranges from 25 to 31 degrees C.     

Temperature-dependent development and parasitism rates of four species of Pteromalidae (Hymenoptera) parasitoids of house fly (Musca domestica) pupae

Parasitoid development, parasitoid-induced host mortality and parasitoid progeny emergence were determined at five constant temperatures for Muscidifurax raptor Girault and Sanders, Muscidifurax zaraptor Kogan and Legner, Spalangia cameroni Perkins and Spalangia endius Walker using pupae of the house fly, Musca domestica L., as hosts. At temperatures of 20, 25, 30 and 35 degrees C the median development times (days from oviposition to adult emergence), respectively, were M. raptor (28.4, 20.7, 14.3, 14.5), M. zaraptor (30.6, 22.8, 14.1, 14.2), S. cameroni (55.6, 35.2, 21.8, 25.0) and S. endius (52.4, 31.5, 16.3, 14.6). All species failed to emerge at 15 degrees C. Using densities of five parasitoids and 100 hosts and a 24 h exposure period, Muscidifurax species oviposited at a greater rate over a wider range of temperatures than Spalangia species. At 15, 20, 25, 30 and 35 degrees C the mean number of pupae killed per parasitoid were, respectively, M. raptor (1.4, 7.4, 10.5, 13.7, 14.1), M. zaraptor (0.0, 3.3, 8.9, 14.4, 15.0), S.cameroni (0.0, 7.8, 11.0, 11.9, 7.4), S.endius (0.6, 4.0, 7.5, 12.0, 11.7), and means of the number of parasitoid progeny per parasitoid were, respectively, M.raptor (0.2, 5.2, 7.9, 11.8, 11.6), M.zaraptor (1.3, 4.4, 8.2, 13.0, 13.7), S.cameroni (0.0, 2.4, 4.7, 5.1, 1.0), S.endius (0.0, 0.9, 3.4, 7.5, 4.9). Development and ovipositional activity in S.cameroni was strongly inhibited at 35 degrees C. The model by Sharpe & DeMichele (1977) was used to describe temperature-dependent development and the number of parasitoid progeny produced per parasitoid at temperatures of 15-30 degrees C in all species.     

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.