Autecology of the tailed jay butterfly Graphium agamemnon (Lepidoptera : Rhopalocera : Papilionidae)

The Tailed Jay Graphium agamemnon is one of the attractive papilionid butterflies that enliven the environment of Visakhapatnam. It occurs throughout the year. It lays eggs singly on young leaves of the mast tree Polyalthia longifolia var. pendula (Annonaceae). The eggs take 3-4 days to hatch. The larvae go through 5 instars over a period of 15-16 days. The pupal period is 13-14 days. The total period from egg to adult emergence spans over 33-36 days. Based on this short life cycle, and larval and pupal development success studied every month, this butterfly can be multivoltine with a minimum of 7-8 broods in a year. Both CI and GR decreased with the age of larva, their average figures being 3.78 and 0.43 respectively. AD values are high (average 92%) and decreased through successive instars. Both ECD and ECI followed a similar pattern with an increase from instar I up to II, then a decrease up to IV and again an increase in instar V and the highest value is with fifth instar. Adults frequently visited flowers (12-35 flowers in a min) spending 1.0 to 3.2 seconds on a flower. The nectar concentration ranged between 16 and 58%. Peak foraging activity mostly fell between 0900-1000 h. The proboscis received pollen in most of the floral species visited, thus satisfying one of the characteristics of butterfly pollination. Being a fast and strong flier it is treated as "high energy" pollinator promoting cross-pollination.     

The occurrence of aristolochic acids in neotropical troidine swallowtails (Lepidoptera: Papilionidae)

Summary. Analysis of individuals of 17 troidine species demonstrated the presence of aristolochic acids in these butterflies in varying concentrations. Although aristolochic acids do not occur in Aristolochia galeata leaves, they were present in Battus polydamas larvae reared on these leaves, and thus may be synthesized by the larvae from chemical precursors in the plant. Received 17 August 1999; accepted 20 January 2000     

Autecology of the common mormon butterfly,Papilio polytes (Lepidoptera : Rhopalocera : Papilionidae)

The adults of the common mormon butterfly Papilio polytes Linn. feed on a variety of floral species. The larval food plants in the study area included Citrus limon and Murraya koenigii both of the family Rutaceae. The eggs are laid singly, and the hatching time is three days. The larvae pass through five instars. The larval growth is directly correlated with the quantity of food consumed. The AD (approximate digestibility) values decreased from first instar to the last, whereas the ECD (efficiency of conversion of digested food) and ECI (efficiency of conversion of ingested food) values increased, thus bearing an inverse relationship with AD. The development time from egg to adult is 28-30, giving 11-12 generations in a year, but with better breeding during August-February. Thus P. polytes is multivoltine.     

Biotransformation of 4-nerolidylcatechol by Heraclides brasiliensis (Lepidoptera: Papilionidae) reduces the toxicity of Piper umbellata (Piperaceae)

Heraclides brasiliensis (Lepidoptera: Papilionidae) larvae feed preferably on Piperaceae, foraging successfully on leaf tissues even though species of this contain high levels of secondary metabolites such as amides and lignans, associated with diverse biological activities including insecticidal properties. Studies examining the metabolism of chemical constituents in Piperaceae by insects are rare. In this study, we characterized the metabolites of 4-nerolidylcatechol (4-NC), the major constituent of Piper umbellata (Piperaceae), and E-2,3-dihydro-3-(3,4-dihydroxyphenyl)farnesoic acid, compounds from fecal material of H. brasiliensis larvae fed a diet containing only P. umbellata leaves. The biotransformed product was also detected in larval and pupal tissues. Moreover, we observed deactivation of the toxicity of P. umbellata leaves against brine shrimp after their metabolism in H. brasiliensis larvae from a LC₅₀ of 523.3 to 3,460.7 μg/mL. This deactivation is closely associated with the biotransformation of 4-NC to E-2,3-dihydro-3-(3,4-dihydroxyphenyl)farnesoic acid, which showed LC₅₀ of 8.0 and >1,000 μg/mL, respectively.     

Aristolochic acid content of South-East Asian troidine swallowtails (Lepidoptera: Papilionidae) and of Aristolochia plant species (Aristolochiaceae)

Summary. Analysis of South-East Asian troidine swallowtails revealed high variability in the content of aristolochic acids among individuals. The presence or absence of these compounds depends on the Aristolochia species available as food plant for the larvae. Only one plant species (Aristolochia philippinensis) contained a high concentration of aristolochic acids, while other species from various localities contained none or only marginal amounts. Whether aristolochic acids have a distinct function in chemical defense of these swallowtails is still an open question. Received 11 December 2000; accepted 4 August 2001.     

Function of the mating plug in Drosophila hibisci Bock

The mating plug in Drosophila hibisci Bock is a firm, gelatinous structure that forms within the female’s uterus during copulation. Two non-mutually exclusive hypotheses for the function of the plug were evaluated. The plug may serve as a nutritional gift that females digest, using the constituents for somatic maintenance or to provision eggs as they mature within the ovaries. Alternatively, the plug may act as a chastity enforcement device by preventing subsequent copulations, and thereby reducing sperm competition. Plug size did not decrease within females over a period of 2 days, and dietary treatment in females did not affect plug size. The extent of ovarian provisioning was also not related to plug size. These results weaken the nutritional gift hypothesis. In contrast, the probability of a second copulation increased sharply with an experimental decrease in plug size. Moreover, females with plugs experimentally reduced in size were courted significantly more and mated significantly faster than females with larger plugs. These results support the chastity enforcement hypothesis. The plug retains the ejaculate and concentrates sperm at the anterior end of the uterus near the apertures of the sperm storage organs. The presence of the plug thus probably facilitates the movement of sperm into storage by retaining sperm at the anterior end of the uterus near the apertures of the sperm storage organs, which may be especially important for D. hibisci, in which sperm length is nearly twofold greater than ventral receptacle length. Matings with newly eclosed virgin females were significantly shorter than with older virgins, and copulations with the younger virgins ended more often without any sperm having yet entered into storage. The effectiveness of the plug in safeguarding a male’s ejaculate may have favoured the evolution of shortened copula durations with young virgins. One fitness advantage of shortened copula duration could be time liberated for the pursuit of further mating opportunities. Received: 12 May 2000 / Revised: 22 September 2000 / Accepted: 15 October 2000     

Florida red bay (Persea borbonia) leaf extracts deter oviposition of a sympatric generalist herbivore, Papilio glaucus (Lepidoptera: Papilionidae)

Summary. The ability to perceive and respond to phytochemicals that reliably indicate poor suitability of a potential host plant confers a selective advantage to ovipositing female swallowtail butterflies. Papilio glaucus females are generalists that nonetheless do not oviposit on red bay (Persea borbonia: Lauraceae). Red bay is toxic to P. glaucus neonates but is commonly found in habitats alongside their principal host plant, Magnolia virginiana, in central Florida. The hypothesis that deterrent compounds present in the leaves of red bay mediate its rejection by P. glaucus was evaluated in our study. Florida populations of P. glaucus did not oviposit on host leaves sprayed with the methanol extract of red bay foliage, although they accepted solvent-treated and untreated tulip tree leaves in 3 choice bioassays. Additionally, tulip tree leaves sprayed with methanolic extracts of red bay also deterred oviposition by P. glaucus females from Ohio, Kentucky and Pennsylvania, although these populations do not naturally encounter red bay. Clearly, deterrent compounds found within this non-host are the basis of its rejection by populations of P. glaucus and such recognition is fundamental to the species, not just a reflection of local adaptations. Received 2 April 1999; accepted 11 June 1999.     

Effectiveness of tiger moth (Lepidoptera, Arctiidae) chemical defenses against an insectivorous bat (Eptesicus fuscus)

Summary. Adult tiger moths exhibit a wide range of palatabilities to the insectivorous big brown bat Eptesicus fuscus. Much of this variation is due to plant allelochemics ingested and sequestered from their larval food. By using a comparative approach involving 15 species from six tribes and two subfamilies of the Arctiidae we have shown that tiger moths feeding on cardiac glycoside-containing plants often contain highly effective natural feeding deterrents. Feeding on pyrrolizidine alkaloid-containing plants is also an effective deterrent to predation by bats but less so than feeding on plants rich in cardiac glycosides. Moths feeding on plants containing iridoid glycosides and/or moths likely to contain biogenic amines were the least deterrent. By manipulating the diet of several tiger moth species we were able to adjust their degree of palatability and link it to the levels of cardiac glycosides or pyrrolizidine alkaloids in their food. We argue that intense selective pressure provided by vertebrate predators including bats has driven the tiger moths to sequester more and more potent deterrents against them and to acquire a suite of morphology characteristics and behaviors that advertise their noxious taste.     

Chemical basis for differential acceptance of two sympatric rutaceous plants by ovipositing females of a swallowtail butterfly, Papilio polytes (Lepidoptera, Papilionidae)

Summary. In the natural habitat of Papilio polytes, a Rutaceae feeder utilizing Toddalia asiatica as a major host plant, some other rutaceous plants such as Murraya paniculata (abundant) and Glycosmis citrifolia (relatively rare) occur sympatrically as potential hosts. Whereas G. citrifolia is occasionally infested in the field, M. paniculata remains entirely unexploited by the butterfly. We thus examined the phytochemical mechanisms that can explain the differential acceptance of the two plants by ovipositing females of P. polytes. The foliage of G. citrifolia was found to readily induce oviposition and females deposited eggs in response to a methanolic extract of the plant. Stimulatory activity-directed fractionation of the extract revealed the presence of two characteristic compounds, trans-4-hydroxy-N-methylproline and 2-C-methylerythronic acid, known to serve as oviposition stimulants for the butterfly. In addition, larvae performed as well or better onG. citrifolia as on T. asiatica. Similar examination of the inhibitory chemical constituents of M. paniculata led to the isolation of an oviposition deterrent. The compound, identified as trigonelline (N-methylpyridine- 3-carboxylic acid), exhibited moderate oviposition deterrency to females. These results combined with our previous data are in agreement with the observed differential utilization of the two plants by P. polytes in the field.     

Male size,mating success,and mating strategy in the mosquitofish Gambusia affinis (Poeciliidae)

Summary In laboratory choice experiments, receptive female western mosquitofish Gambusia affinis affinis (deprived of contact with males 30 days) preferred the larger of two males. When two males differing in size were placed with a receptive female, the larger was generally able to monopolize access to her, but not when the female was not receptive. In other experiments, a single male was placed with a receptive or non-receptive female. These experiments showed size-correlated differences in male mating behavior. Smaller males were more likely to chase non-receptive females and attempt to inseminate them forcibly, while larger males continued to court females even if they were non-receptive. Even when the female was receptive, there was some evidence of size-correlated differences in levels of courtship and chasing.     

Alternative mating tactics in males of Polistes dominulus (Hymenoptera: Vespidae)

Summary In summer, males of Polistes dominulus form large aggregations at sunny landmarks. We identified two size-correlated behavioural categories: residents (R) and transient (T). R males, which constitute 20%–25% of the total population, are larger than T males, territorial, aggressive, and more site-faithful, while T males range more widely, are non-aggressive, and show little site tenacity. Field and laboratory data suggest that R males have an advantage in mating, particularly if they engage in frequent flights while on their territories. These alternative mating tactics within the same population are combined with behavioural flexibility in some individuals, which switch from one option to the other.     

Lifetime mating success in a natural population of the damselfly,Enallagma hageni (Walsh) (Odonata: Coenagrionidae)

Summary Variance in lifetime mating success was measured for individuals of a population of Enallagma hageni, a non-territorial damselfly in northern Michigan. E. hageni is an explosive breeder with scramble competition for mates. Highly skewed operational sex ratios resulted in intense male-male competition which took the form of interference with tandem pairs. 41% of the males failed to mate in their lifetime as opposed to only 3.6% mating failure in females. The effect on mating success of size, age, longevity, and time spent at the breeding site were investigated. Intermediate sized males obtained the most matings, and male lifetime mating success was highly correlated with longevity.     

The cause of size-assortative mating in the leaf beetle Trirhabda canadensis (Coleoptera : Chrysomelidae)

Summary Examples of positive assortative mating by body size are abundant but its causes remain controversial. I show that size-assortative mating occurs in the chrysomelid beetle Trirhabda canadensis and I test a series of alternative hypotheses to explain how this mating pattern comes about. Results suggest that assortative mating in this beetle is due to the greater ease with which size-matched pairs can achieve intromission, and not due to size-biased skews in the availability of mates or mate choice favoring large individuals. There was no correlation between male and female elytron length (a measure of body size) at the initiation of courtship, but pairs assorted positively by size at the onset of intromission. Moreover, in the laboratory, there was a negative correlation between male and female size for pairs engaged in courtship that terminated without mating. Assortative mating was not associated with a large-male mating advantage and there was no evidence of female choice of large males. Nor was there unequivocal evidence for male choice of large females; although mating females were slightly larger and considerably heavier than solitary females, males did not differ in the frequency with which they rejected large and small females. Assortative mating in T. canadensis appeared to be caused by the lower ability of mismatched pairs to achieve intromission after an encounter, both when males were larger and when they were smaller than the female.     

Tropane and pyrrolizidine alkaloids in the ithomiinesPlacidula euryanassa andMiraleria cymothoe (Lepidoptera: Nymphalidae)

Summary Larvae of the ithomiine butterflyPlacidula euryanassa sequester tropane alkaloids (TAs) from the host plantBrugmansia suaveolens and pass them through the pupae to freshly emerged adults. Wild caught adults also show in their tissues, variable amounts of pyrrolidizine alkaloids (PAs), probably sequestered from variable plant sources and subject to dynamics of incorporation, accumulation and utilization of PAs by ithomiine butterflies. The ratio TAs/PAs is also variable between different populations.Miraleria cymothoe, another ithomiine that feeds onB. suaveolens as larvae, does not sequester TAs from the host-plant, but sequesters PAs from plant sources visited by the adult butterflies. The main alkaloid found in both butterflies is lycopsamine, which also is the principal PA found in all genera of Ithomiinae.     

Evidence for division of labor in the social caterpillar Eucheira socialis (Lepidoptera: Pieridae)

The caterpillars of Eucheira socialis westwoodi cooperatively spin and maintain a hollow silken nest and an elaborate network of silken foraging trails on their host plant, madrone (Arbutus spp.: Ericaceae). Nests typically contain several hundred larvae. Two populations are known to harbor a sex ratio distorter. The primary sex ratio in these two populations for four generations has been exceedingly male biased (64–79% male). Lepidoptera larvae are easily sexed using external morphology, allowing us to uniquely mark male and female larvae and to assemble larval groups of particular sex ratios. We report here the results of experiments on sex-specific larval behavior and physiology and the effect of colony sex ratio on individual behavior. We found that male larvae spent more time spinning silk on the nest and less time feeding than female larvae. Males were the first to emerge from the nest and the first to venture out along trails to feed. Male-biased nests had a significantly greater amount of silk deposited on their surfaces than female-biased nests. In the field, male-biased nests produced heavier male and female pupae than female-biased nests. Male and female larvae in 75% male nests became active earlier than males and females in other sex ratio treatments. Received: 11 September 1998 / Received in revised form: 24 February 1999 / Accepted: 27 March 1999     

Toxicity of nonhost phototoxins to parsnip webworms(Lepidoptera: Oecophoridae)

Summary The parsnip webworm,Depressaria pastinacella (Lepidoptera: Oecophoridae), feeds exclusively on apiaceous hostplants containing furanocoumarins, compounds capable of oxygen-dependent and oxygen-independend photosensitization. Despite high titers of antioxidant enzymes relative to other herbivorous insects, webworms cannot tolerate nonhost photosensitizers such as alpha-terthienyl or beta-carboline alkaloids at dietary concentrations of 0.01% or less. Tolerance of skimmianine, a furano-quinoline alkaloid, may be due to its structural resemblance to furanocoumarins, which are metabolized by cytochrome P450 monooxygenases in this species.     

Determinants of male mating success in the red bishop (Euplectes orix)

We studied sexual selection in the red bishop, Euplectes orix, a colonial, polygynous weaverbird widely distributed over sub-Saharan Africa. Male reproductive success measured in terms of the number of nests accepted by females and the number of eggs and nestlings in all the nests on a male's territory varied considerably. The standardized variance (variance/mean²) in male reproductive success ranged from 0.505 to 1.737 in different years, indicating a high potential for sexual selection in this species. An analysis of genetic parentage for 432 nestlings by non-radioactive, multilocus DNA fingerprinting confirmed that male reproductive success (number of young sired on the territory) in this species can be reliably estimated by the measures introduced above. In all 4 study years there was a strong positive correlation between male mating success and the total number of nests that males built in their territories. The number of nests built can be partitioned into the number of weeks a male held a territory and his nest-building performance. Both factors exert a significant positive effect on male mating success and in combination explained between 53.3 and 86.3% of the variation in male reproductive success. Male morphological characters were found to be of no importance. Males that established a territory in the following season built more nests and held their territories for longer than males that did not establish a territory in the following season, suggesting that these measures might be indicators of male condition and quality. Male nest-building performance (number of nests built per week) seems to be unrelated to male condition or quality. Received: 8 January 1999 / Received in revised form: 7 June 1999 / Accepted: 13 June 1999     

Male mating success and paternal care in Nicrophorus vespilloides (Coleoptera: Silphidae)

Summary If no female is present, male burying beetles Nicrophorus vespilloides (Coleoptera: Silphidae) co-operate in the burial of a corpse. Once a female has arrived, the males fight with one another. The defeated male stays near the corpse and to copulate with the female. Laboratory experiments using sterilised males showed that the defeated male was able sometimes to father some of the offspring raised on the corpse. Male N. vespilloides almost always participate in defence and feeding of the brood. This is not affected by the size of the male. Males quickly leave or are driven from 5 g corpses. Males feed the larvae as often as females do, and larvae raised by males alone are not significantly different in weight from larvae raised by females alone or by both parents. Males which cared for a succession of broods in the laboratory did not differ significantly in median lifespan from males which were removed from their corpses after eggs had been laid. Non-caring males weighed significantly more than caring males over a sequence of corpses, but the caring males did not differ significantly in weight from non-breeding controls.     

Sexual selection and mating system in Zorotypus gurneyi Choe (Insecta: Zoraptera)

Body size is clearly an important factor influencing the outcome of agonistic contests, but is often weakly correlated with dominance ranks in Zorotypus gurneyi Choe (Insecta: Zoraptera). The study of the development and dynamics of dominance relations using artificially constructed colonies show that age, or tenure within the colony, is the prime determinant of dominance among males. Dominance hierarchies become relatively stable within 2 or 3 days and males that emerge later normally begin at the bottom of the hierarchy regardless of size. Males interact much more frequently when they are simultaneously introduced to each other than when they are allowed to emerge at different times. In the latter case, males that emerge late appear to recognize relative dominance of older males and avoid direct contests. Considering the high correlation between dominance rank and mating success, there is a strong selective advantage to males that emerge earlier and such pressure of sexual selection may be responsible for the difference in life history strategies between Z. gurneyi and its sympatric congener, Z. barberi Gurney, in central Panama.