Graphium agamemnon Linnaeus (Lepidoptera: Papilionidae), a pest of soursop (Annona muricata Linnaeus), in Vietnam: Biology and a novel method of control

Few butterflies are pests of economic significance, but some may be locally destructive, such as the papilionid Graphium agamemnon Linnaeus, which is known to feed on the commercially important soursop (Annona muricata Linnaeus) in Vietnam. This paper documents the life history and ecology of G. agamemnon and investigates commonly used control measures in south‐east Vietnam. A novel method of controlling G. agamemnon infestation is described. If soursop basal rootstock (Annona glabra Linnaeus) is encouraged to shoot during times of peak butterfly activity, ovipositing G. agamemnon females are preferentially attracted to the new growth where the resulting early stages may be easily detected and removed by hand.     

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.     

New evidence on the systematic and phylogenetic position of Parides burchellanus (Lepidoptera: Papilionidae)

Parides burchellanus is considered a rare and threatened swallowtail species restricted to central Brazil. It shows considerable morphological similarity to Parides panthonus jaguarae, with which it shares both geographical range and larval host plant. At present, P. burchellanus and P. panthonus are believed to be distinct species, based on minor differences in male genitalia. In this study, the phylogenetic and systematic position of P. burchellanus in relation to three subspecies of P. panthonus (P. p. jaguarae, P. p. lysimachus and P. p. aglaope) was evaluated using molecular evidence: the complete sequence of the mtDNA genes COI and COII and of the nuclear gene EF‐1α (c. 3300 bp). In addition, the informativeness of the ‘barcode’ region next to the 5′ end of COI (c. 650 bp) was evaluated for delimiting these taxa. Individual analysis by neighbour‐joining, using Kimura 2‐parameter distance model, and by maximum parsimony showed that P. p. jaguarae + P. p. lysimachus + P. p. aglaope + P. burchellanus form a strongly supported monophyletic clade, and all molecular regions consistently recovered P. p. jaguarae and P. burchellanus as sister species. The genetic divergence among the subspecies of P. panthonus and P. burchellanus is equivalent to the divergence among conspecifics of other species of Parides, and smaller than the interspecific divergence among different sister species of this genus. The results support the proposal that P. p. jaguarae and P. burchellanus are likely to be synonymous, and suggest that P. burchellanus can be considered conspecific with P. panthonus. The taxonomic classification of P. burchellanus should be revised on the basis of the molecular data.     

Behaviour of roach (Rutilus rutilus L.) altered by Ligula intestinalis (Cestoda: Pseudophyllidea): a field demonstration

1. We studied the influence of a cestode parasite, the tapeworm Ligula intestinalis (L.) on roach ( Rutilus rutilus L.) spatial occupancy in a French reservoir (Lake Pareloup, South-west of France).
2. Fish host age, habitat use and parasite occurrence and abundance were determined during a 1 year cycle using monthly gill-net catches. Multivariate analysis [generalized linear models (GLIM)], revealed significant relationships ( P  < 0.05) between roach age, its spatial occupancy and parasite occurrence and abundance.
3. Three-year-old roach were found to be heavily parasitized and their location toward the bank was significantly linked to parasite occurrence and abundance. Parasitized fish, considering both parasite occurrence and abundance, tended to occur close to the bank between July and December. On the contrary, between January and June no significant relationship was found.
4. These behavioural changes induced by the parasite may increase piscivorous bird encounter rate and predation efficiency on parasitized roach and therefore facilitate completion of the parasite's life cycle.     

The influence of a shortage of hosts on the specificity of oviposition behaviour in species of Dacus (Diptera, Tephritidae)

ABSTRACT. When deprived of their preferred hosts for up to 16 days prior to behaviour assays, three relatively specialized tephritid flies, D. cacuminatus (Hering), D. cucumis French and D. jarvisi (Tryon), retain strong preferences for their particular hosts. Moreover, in the absence of the preferred hosts, D. cacuminatus , and to a lesser extent D. cucumis , will not accept fruits that are outside their normal host range. D. jarvisi readily accepts other fruits in the absence of its preferred host, Planchonia careya , but in common with the other two shows no increase in the number of eggs laid with increasing periods of deprivation. None of these species accumulate oocytes when deprived of fruits.
In contrast, the highly polyphagous D. tryoni Frogg. lays more eggs and oviposits readily into a previously unacceptable fruit after 4 days without access to hosts. This change in behaviour is associated with a marked increase in the number of mature eggs carried by females.
These results suggest an important difference between the generalist and more specialized species in the physiological control of oocyte maturation. In D. tryoni there appears to be no inhibition of oocyte development once the primary follicle has matured, whereas in the other species each ovariole contains no more than one mature egg.
In D. tryoni , the change in behaviour which accompanies this increase in egg load, may influence its propensity to accept unusual fruits, and may help to explain its rapid acceptance of most cultivated fruits introduced to Australia.     

豚草生育期和被利用状态对广聚萤叶甲产卵选择的影响

【背景】植食性昆虫对产卵场所的选择行为受多种因素的影响,其中寄主植物的发育阶段和被同种其他个体利用程度是影响其产卵选择的重要因素之一。【方法】采用野外罩笼的选择性试验,观察了广聚萤叶甲成虫对不同生长发育阶段、不同被取食程度、不同落卵量的豚草植株的产卵选择行为,分析产卵选择频次,用Cox比例风险模型拟合识别期以估计产卵选择偏好性。【结果】开花期豚草被选择产卵的风险(瞬间概率)仅为生长期的16.5%,苗期豚草被选择产卵的风险为生长期的34.1%;成虫对3种被食程度豚草的产卵选择频次之间存在显著差异(n=20,P<0.01),在3级被食程度豚草上产卵次数最多(12次),而在5级被食豚草上产卵次数最少(2次),表明成虫明显偏好将卵产于已被中等程度取食的植株上;成虫对不同落卵量植株(0、4、8个.株-1)未表现出显著的产卵偏好,对不同密度(高、低)斑块的豚草的产卵选择亦无显著差异。【结论与意义】豚草生育期和被取食程度影响广聚萤叶甲雌虫的产卵选择,而植株上的落卵量和豚草密度对产卵选择没有影响。     

Comparison of oviposition behavior of a solitary and a gregarious parasitoid (Hymenoptera: Mymaridae)

Anaphes victus Huber andAnaphes listronoti Huber (Hymenoptera: Mymaridae) are respectively solitary and gregarious egg parasitoids of the carrot weevil,Listronotus oregonensis (LeConte) (Coleoptera: Curculionidae). We made detailed ethograms of the oviposition behavior on unparasitized and parasitized hosts for the two species. We then compared the behavior of virgin and mated females for the oviposition of male and female progenies. The two species did not always oviposit after insertion of the ovipositor, but these punctures without oviposition could be readily differentiated from oviposition.A. victus oviposited only once by puncture, whileA. listronoti deposited one to three eggs during the same sequence. The variability of the duration of the various components was generally lower for a given female than between females. Two components, the abdominal vibrations and the pause, were significantly shorter in ovipositions that resulted in male progency for the two species. However, an important overlap in duration prevents using these differences to sex the progeny at oviposition. Virgin females of both species, although capable of producing only males, exhibited both behaviors. Parasitized hosts were recognized through internal and external markings that were used in host discrimination.     

Oviposition on host plants by a tropical forest grasshopper (Microtylopteryx hebardi: Acrididae)

Abstract. 1. The tropical forest grasshopper Microtylopteryx hebardi Rehn (Orthoptera, Acrididae, Ommatolampinae) oviposits inside leaves or stems of its host plants. This is the first known instance of oviposition inside living plant tissue (rather than in the ground, on top of leaves, or in detritus) for a tropical forest acridid.
2. In understorey secondary forest at La Selva, Costa Rica, M.hebardi females fed on fifteen species of host plants, but oviposited in only four of these.
3. In laboratory choice tests, females showed selectivity for oviposition host plants similar to that inferred from field observations.
4. Plant volume was a better predictor of the number of eggs laid per stem than variables that indicated leaf or stem characteristics of individual plants.
5. Unlike the ovipositor of acridids laying eggs in the ground, that of M.hebardi is equipped with toothed, sclerotized edges, probably used to bore into plant tissue.
6. Endophytic oviposition in M.hebardi may be related to finding host plants in a habitat that is complex, heterogeneous, and rich in plant species.     

广聚萤叶甲成虫取食经历对后续 产卵寄主选择的影响

【背景】取食经历对植食性昆虫的寄主选择行为具有较大影响,影响天敌昆虫寄主专一性测定的设计和结果解释。【方法】采用选择性试验,观察了入侵豚草的重要天敌——广聚萤叶甲成虫羽化后取食不同植物对其后续产卵寄主选择的影响。【结果】与取食豚草的试虫相比,有取食三裂叶豚草、苍耳或菊芋经历的成虫选择苍耳产卵的频次增加,不再对豚草表现出明显的选择偏好性。对产卵识别期的Cox模型分析结果表明,成虫早期取食不同植物,对后续产卵选择有显著影响,成虫羽化后如果先取食豚草或三裂叶豚草,则选择苍耳产卵的倾向显著低于豚草;但如果先取食苍耳、菊芋和农家向日葵,则选择苍耳产卵的倾向与豚草无显著差异。【结论与意义】由此推测,广聚萤叶甲初羽化成虫取食的植物对其后续产卵选择具有较大影响,因而在寄主专一性测定中应关注测试前饲喂的植物种类。     

Effect of sorghum resistant to Stenodiplosis sorghicola (Coquillett) (Diptera: Cecidomyiidae) on oviposition and development of Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae)

Abstract Helicoverpa armigera oviposition preference for, and larval development on sorghum hybrids with differing resistance to sorghum midge, Stenodiplosis sorghicola , were investigated. When H. armigera larvae were fed seed of resistant and susceptible hybrids in the laboratory there were no differences in larval and pupal sizes or the rate of development. The same result was recorded when larvae fed on panicles on plants in a glasshouse. On some sampling occasions, significantly more eggs were laid on panicles of resistant hybrids in the field. This occurred when plants were in plots and also in a mixed planting. Midge-resistance status did not affect levels of egg parasitism. In a field study using recombinant inbred lines between a midge-resistant and a midge-susceptible line, no relationship was found between level of resistance and oviposition of H. armigera . We conclude that, although midge-resistant hybrids are sometimes preferred for oviposition by H. armigera, the resistance per se does not determine this preference. Egg survival, larval survival, development and resultant damage are not significantly affected by the midge-resistance status of the host.     

Effect of host deprivation on egg quality,egg load,and oviposition in a solitary parasitoid,Chetogena edwardsii (Diptera: Tachinidae)

Solitary parasitoids are limited to laying one egg per host because larvae compete within hosts. If host encounter rate is low, females should not increase the number of eggs/host in response. The tachinid fly, Chetogena edwardsii,was used to evaluate the effect of host deprivation on egg accumulation, oviposition behavior, and egg quality in a solitary parasitoid. Females deprived of hosts for 2– 7 days accumulate about 1 day's supply of eggs. Egg output of deprived females once hosts are restored does not differ from that of control females. Deprived females retain one egg in the uterus where it undergoes embryogenesis. Maggots emerging from retained eggs are more likely to survive in hosts molting in 40 h or less after receipt of an egg than are maggots emerging from eggs fertilized shortly before oviposition. Egg retention is a consequence of host deprivation that permits females to broaden the range of hosts they can exploit to include soon-to-molt hosts and possibly multiply parasitized hosts.     

Effects of physiological condition and experience on oviposition behaviour of Trichogramma australicum Girault (Hymenoptera: Trichogrammatidae) on eggs of Helicoverpa armigera Hübner (Lepidoptera: Noctuidae)

We produced an ethogram for and investigated effects of physiological condition and experience on oviposition behaviour for Trichogramma australicum on Helicoverpa armigera eggs. The data, coupled with the ethogram, have enabled the establishment of standard behavioural categories which can be used to assess acceptability of H. armigera eggs to T. australicum reared in vitro or in eggs of other species. Physiological and experiential conditions were investigated using females of three different ages (0–24, 24–48 and 48–72 h old), two types of sexual states (mated and unmated) and two types of ovipositional experience (naive and experienced). Durations of host finding, host examination, and post-ovipositional re-examination were reduced by oviposition experience and were shorter in younger females (0–48 h old). Oviposition experience reduces the duration of the drilling and percentage of females host-feeding. Being mated and young reduces the host examination time in subsequent oviposition bouts. Only ovipositionally inexperienced females host-feed before oviposition and host-fed more frequently than experienced individuals after oviposition. Host-feeding needs consideration for optimal culture.     

Effects of cultivar on oviposition preference, larval feeding and development time of diamondback moth, Plutella xylostella (L.) (Lepidoptera: Plutellidae), on some Brassica oleracea vegetables in Victoria

Abstract We studied oviposition preference of Plutella xylostella for four cultivars of cabbage, broccoli and cauliflower in the field. There were no differences in the number of eggs found on the various cultivars of broccoli or cauliflower. Significantly more eggs were laid on cultivar Savoy King than any of the other cabbage cultivars tested. We then compared the development time and feeding damage of larvae reared on Savoy King and Green Coronet cabbages. Larvae developed more rapidly and fed more and for longer on Green Coronet than Savoy King. Thus, while Savoy King is more attractive to oviposition in the field, net impacts on the crop may be lessened to some degree through lower feeding proficiency of the larvae on that cultivar.     

高岭土颗粒涂布对中国梨喀木虱寄主选择、产卵的影响及应用效果

为探索可替代化学药剂防治中国梨喀木虱的措施,开展了高岭土颗粒涂布对其寄主选择、产卵影响及应用效果的研究。分别采用3.5%高岭土水溶液、清水对梨休眠枝和当年生枝叶进行喷雾涂布处理,于24、48、72、96 h监测选择条件下成虫分布情况,于96 h记录选择和非选择条件下各枝条上的卵量。采用高岭土水溶液对梨苗均匀喷雾涂布,通过被害叶率、单叶若虫量、虫龄指数、为害指数对应用效果进行评价。研究结果显示,在选择条件下,经高岭土涂布的枝条上成虫数量明显少于对照,高岭土涂布的枝条上冬型成虫数量在96 h时最低(0.5头/枝),96 h时着卵量(1.83头/枝)显著低于对照(30.83头/枝);高岭土涂布的枝叶上夏型成虫在72 h时最低(1.17头/枝),96 h时着卵量(12.33头/枝)显著低于对照(242.5头/枝);在非选择条件下,冬型和夏型成虫在高岭土涂布的枝条上着卵量(2.83和15.17头/枝)分别显著低于对照(25.17和170.42头/枝);供试植株在接虫45 d后为害最严重,高岭土涂布的植株被害叶率(54.17%)、单叶若虫量(2.29头/叶)、虫龄指数(3.79)、为害指数(0.41)分别显著低于为对照植株(98.61%、11.72头/叶、4.13、0.95)。上述结果表明,高岭土涂布对中国梨喀木虱成虫的寄主选择具有驱避效果,对雌成虫的产卵具有抑制作用;高岭土涂布可限制中国梨喀木虱种群增长,降低危害水平。研究结果为中国梨喀木虱的综合治理提供了现实依据。     

Poly modal emergence of the tiger swallowtail, Papilio glaucus (Lepidoptera: Papilionidae): source of a false second generation in central New York State

Abstract. 1. The tiger swallowtail butterfly, Papilio glaucus (L.) (Lepidoptera: Papilionidae), is commonly assumed to have a bivoltine life cycle in central New York State and other parts of the northeastern United States. The seasonal abundance of adults in Tompkins County, New York, shows a bimodal pattern in some years, and a skewed unimodal pattern in others. We studied the incidence of diapause, the potential for successful development of two generations in the field, and the dates of adult emergence from overwintering pupae to determine whether these seasonal patterns are the result of bivoltinism.
2. Insects from Tompkins County, reared as larvae under 16L:8D photoperiod, uniformly enter pupal diapause.
3. Comparison of the heat units required for successful development of two generations with the heat units available in the field during 11 years shows that emergence of non-diapausing individuals would occur too late in the summer to contribute to the second peak of butterfly abundance. Offspring of non-diapausing individuals could only rarely complete development before the end of the growth season.
4. Diapausing offspring of butterflies collected in early June and late July differed sufficiently in dates of emergence to account for the field pattern of seasonal abundance.
5. Electrophoretic studies revealed no evidence of genetic isolation between early and late emerging butterflies.
6. These results suggest that polymodal emergence of diapausing insects can masquerade as a bivoltine life cycle.     

Subsocial behaviour in Oniticellus cinctus (Coleoptera, Scarabaeidae): effect of the brood on parental care and oviposition

ABSTRACT. In Oniticellus cinctus (F.) the nest chambers each contain about twenty brood balls. Females enlarge the brood balls during the egg and larval stages and remain in the chamber for the whole period of brood development (1 month); they then make a new nest after 1 week. The presence of the brood releases parental care and ensures that the mother remains in the nest: she repairs defects in the brood balls and the nest, and expels other O. cinctus females. A new ball is formed around a naked O. cinctus larva, but larvae of other species are killed. In addition, the brood inhibits oviposition: removal (or addition) of brood balls stimulates (or inhibits) egg laying. In inhibited ovarioles, existing follicles are resorbed and production of new ones ceases. Control of clutch size by the brood is an adaptation to the nest structure and life history of O. cinctus. It may have an important role in the reproductive strategy of other insects with parental care.     

Ecological significance of puddling as a behavioural phenomenon in the life history of butterfly Papilio polytes Linn. (Lepidoptera: Papilionidae)

Puddling in butterflies is an essential physiological phenomenon, by which they gain mineral nutrients required for their reproductive success and flight. The preference given to the puddling by butterflies has been analysed by two hypotheses: butterflies puddle when there is a scarcity of nutritional resource in one hypothesis and competition with other males to increase their probability of mating in the other. In the present investigation, an attempt has been made to verify these hypotheses through experimental evidences in the butterfly Papilio polytes, in which only males are known to puddle. Our investigation revealed the significance of puddling as an alternative source of nutrition in the absence of nectar. However, increase in the number of males in a given vicinity with constant number of females enhances the puddling activity in males. This shows the importance of puddling as a distinct competitive behaviour by which male improves their chances of mating. Additionally, in both cases the puddling is drastically regulated by the environmental temperature. Males prefer to puddle at a temperature ranging from 26 °C to 28 °C, which is considered as optimum for puddling activity. Further, the fecundity and longevity of imago are completely unaffected by puddling activity. All these results are in good agreement with the existing hypotheses and show the behavioural significance of puddling in butterflies.     

The effect of ejaculate mass on female reproductive output in the european swallowtail butterfly,Papilio machaon (L.) (Lepidoptera: Papilionidae)

Previously, we showed that virgin males of Papilio machaondeliver ejaculates that are twice as big as any ejaculates they transfer at later matings. Here, we investigate the consequences of these two size classes of ejaculates on female reproductive output and demonstrate that females that received one small ejaculate laid as many eggs, fertilized the same proportion of eggs, and lived as long as those that had received one big ejaculate. However, females that received big ejaculates laid heavier eggs, but only between the twelfth and the fifteenth days of egg-laying. We conclude that male-derived nutrients appear to have a limited effect on female reproductive output in P. machaonand that the large size of ejaculates delivered by male butterflies are determined primarily by selection on males to produce longer refractory periods in females.     

Calibration and evaluation of field cage for oviposition study with Aedes (Stegomyia) aegypti female (L.) (Diptera: Culicidae)

Differences among results gathered from insect behavior studies conducted in laboratory and field situations are due to ambient variables that differ greatly between both environments. In laboratory studies the environmental conditions can be controlled whereas in field temperature, humidity and air velocity vary uncontrollably. The objective of this study was to calibrate and evaluate an experimental area (field cage) (14 x 7 x 3.5 m) subdivided into eight test cages (2.5 x 2.5 x 2 m) for use in behavioral oviposition tests of Aedes aegypti (L.) mosquitoes for developing a new methodology to assess attractants and oviposition traps. Test cage calibration involved: (1) minimal experiment duration tests; (2) optimal female release number per traps test and (3) trap placement tests. All tests used gravid A. aegypti females; 3-4 days post blood meal and the sticky trap MosquiTRAP to catch adults. Ninety percent of the females released were recaptured 2h after the beginning of the experiment, and this allowed up to 32 test repetitions/day to be conducted in the field cage. The minimum number of females necessary to conduct statistical analyses was 20 females/trap/test per cage. No significant difference was found in the behavioral response of gravid females to four different trap positions within test cages. Field trapping results with attractant were similar to those in the field cage. Therefore, the field cage could replace field trapping for evaluating at least mosquito traps and oviposition attractants for A. aegypti.