Immature stages of the butterfly Magneuptychia libye (L.) (Lepidoptera: Nymphalidae, Satyrinae)

The immature stages of the butterfly Magneuptychia libye (L.) are described, and their morphology is compared with other Neotropical Satyrinae.     

Homologies in the colour patterns of the genus Heliconius (Lepidoptera: Nymphalidae)

The colour patterns of Heliconius butterflies are composed from a relatively simple set of pattern elements whose homologues are recognizable throughout the genus. Although Heliconius colour patterns look quite different from those of most nymphalids, these pattern elements are seen to derive from the generalized nymphalid groundplan. The differences arise primarily from the loss or positional shift of certain pattern elements, a high degree of fusion between individual pattern elements, and, in the forewing, asymmetries of the pattern elements relative to the wing-cell midline. The scheme of homologies we present is consistent with what is currently known about the comparative morphology and developmental physiology of colour pattern formation in Lepidoptera, and provides a framework for the interpretation of developmental, evolutionary and genetic studies in Heliconius.     

Single cell responses in female Pieris brassicae (Lepidoptera: Pieridae) to plant volatiles and conspecific egg odours

Action potentials from individual olfactory cells and receptor potentials were recorded from antennae of female cabbage white butterflies, Pieris brassicae, using a new ‘surface-contact’ recording technique. Stimuli used were plant volatiles and conspecific egg odours. Most cells were activated by some stimuli and inhibited by others. Hence, the ‘odour spectra’ of most cells included activating as well as inhibiting substances. In addition, the ‘response profile’ of a given stimulus contained both positively and negatively responding cells.Cluster analysis of the odour spectra revealed the existence of moderately separated clusters of olfactory receptors. No groups of cells specifically tuned to odours of eggs or cabbage leaves were found. Analysis of the response profiles did not show a clear clustering of stimuli. The results suggested that the membranes of different receptor cells contain several types of acceptor sites in various proportions.No correlation between EAG and receptor potential amplitude or spike frequency was found. It was concluded that the EAG may be useful in comparing the relative receptor sensitivities only when chemically related compounds are involved.     

Evidence of protease in the saliva of the butterfly Heliconius melpomene (L.) (Nymphalidae, Lepidoptera)

Butterflies of the genus Heliconius are well known for their peculiar habits of utilizing pollen as a source of amino acids. Saliva plays a major role in the process of extracting amino acids and proteins from the pollen grains. In this investigation, we obtained samples of saliva from adult Heliconius melpomene by placing pumpkin pollen or fine glass-beads on the proboscis, which stimulates the butterflies to release saliva. Proteolytic activity was determined in the saliva by an insoluble protein-dye that turns blue when cleaved by proteases. Its extinction value was measured with a spectrophotometer at 595 nm. Both the saliva sampled with pollen and the saliva obtained from inert glass-beads exhibit proteolytic activity demonstrating that the saliva contains proteases. The proteolytic activity of the pollen/saliva samples was higher than that of the glass-bead/saliva samples, which we attribute to the stimulating effects of pollen, such as taste, smell, and texture, and not to proteases which might have been liberated from the pollen. This is indicated by the fact that pollen samples without saliva showed only a negligible indication for proteolytic activity. In general, females exhibit higher proteolytic activities than males, presumably due to their greater amino acid investment in reproduction. We present here first evidence for the existence of proteases in the saliva of a butterfly species and suggest that these enzymes are crucial for the use of amino acids and proteins from pollen in Heliconius butterflies.     

Olfactory sensitivity to amino acids in the juvenile stages of the European eel Anguilla anguilla (L.)

Scanning electron micrograph observations of the olfactory mucosa from both unpigmented glass eel(GE)andpigmentedelvers(EL)of the European eel, Anguilla anguilla (L.), revealed the presence of various cell types; amongst these, the ciliated and microvillous ones are likely to possess a chcmosensory function. Recording of underwater electro-olfactograms (EOGs) showed that various amino acids (glycine, L-alanine, L-valine, L-leucine, L-asparagine, L-glutamine and L-methionine) are effective stimulants for the olfactory mucosa. Dose response curves of stimulus concentrations v. EOG amplitudesfit regression linesat both GE and EL stages. Leucine was more stimulatory at the GE than at the EL stage. The stimulatory effect of the other six amino acids tested was similar at both developmental stages. The possible role of olfactory sensitivity in animal behaviour at different developmental stages is discussed.     

Olfactory Learning of Fruit Odors in the Eastern Yellow Jacket, Vespula maculifrons (Hymenoptera: Vespidae)

Food-seaching workers of eastern yellow jackets, Vespula maculifrons, are attracted by the natural odors of a wide variety of succulent fruits; particularly effective was pear. The only part of a fruit that repelled was the leathery epicarp of oranges. After rewarding with sugar water, odors of six fruits, including the pulpy mesocarp of oranges and, in addition, the leaves of catmint Nepeta cataria, all become highly attractive. To learn the distinctive odors of any one of three fruits (pear, apple, quince), nondiscrimination training with a rewarded fruit was sufficient for the subsequent olfactory preference of the training fruit over the control fruit. In the other cases [banana, hawthorn (Crataegus crus-galli), grape] simultaneous discrimination training with a rewarded and an unrewarded fruit was necessary and effective for obtaining differential responses to the odors of the training fruits. As far as current evidence goes, olfactory learning plays similar roles in the fruit foraging of this wasp and in the nectar foraging of the honey bee (Apis mellifera).     

Visual mate-searching behaviour in the evening brown butterfly, Melanitis leda (L.) (Lepidoptera: Nymphalidae)

Abstract As in most animals, male butterflies are generally the more proactive sex with respect to seeking out mating opportunities. In most cases, males conduct their search sometime between mid-morning and mid-afternoon, but a few species are active only before dawn or after sunset. These crepuscular species offer a good opportunity to study how males deal with markedly different visual and thermal conditions. Here, I present data from a 5-month behavioural study of male Melanitis leda (Nymphalidae) at a mate-encounter site in tropical Australia. Males of this species defended perching locations along a forest edge in a similar manner to other diurnally active territorial nymphalids. They generally arrived at these sites after sunset and arrived earlier on evenings that darkened earlier. Actively mate-locating males were only seen at the site during a 25−35 min evening period, during which ambient light levels ranged between 50 and 2600 lux. Only 27% of marked territory residents were recorded again at their location of capture, but fidelity in this 'resighted' group ranged up to 23 days. A sample of males, captured under ambient temperatures of 24.0−27.2°C, maintained a mean thoracic excess of 8.25 ± 0.73°C, but did not appear to 'shiver' in the manner of other crepuscular species. Males courted conspecific females and one mating was observed. This species is an excellent candidate for further research into the evolution of mating tactics in crepuscular butterflies.     

Possible Age-Dependent Variation in Egg-Loaded Host Selectivity of the Pierid Butterfly, Anthocharis scolymus (Lepidoptera: Pieridae): A Field Observation

To understand better the effects of age on host selection through conspecific egg detection by Anthocharis scolymus females, field observations were performed at a graveyard where the insect population was isolated and the host plant was limited to the cruciferous plant, Turritis glabra. We chased females and recorded their oviposition behaviors and the conditions of plants which females approached. Older females tended to avoid ovipositing on egg-loaded host plants and selectively laid eggs on nonloaded host plants. This result was not confounded by other factors such as seasonality, air temperature, plant height, plant phenological stage, surrounding vegetation, host plant density, and extent of plant damage. We discuss the possibility that females make an oviposition decision on how to lay their limited number of eggs during their remaining lifetime.     

Sexual Isolation Between Two Sympatric Types of the Butterfly Eurema hecabe (L.)

In Japan, the butterfly Eurema hecabe (L.) consists of two separate types. These types differ in wing fringe color (brown or yellow), which is linked with expression of seasonal wing morphs and with ability for host-plant use. It was recently found that these two types occur together on the subtropical Okinawa-jima Island (one of the Ryukyu Islands). In the present work we studied whether sexual isolation may exist between these sympatric types of E. hecabe on Okinawa-jima, using caged conditions. When living females (days 0, 1, and 7) were presented to males of the same or different type, females of the brown type received males of the same type and responded with a refusal posture to those of the yellow type. Females of the yellow type received males of the same type at all ages tested. However, when they were exposed to males of the brown type, the proportion of females showing a refusal posture changed with female age. The proportion was high at day 7 but relatively low at day 0 or day 1. When model females of the brown and yellow types were simultaneously presented to males of each type, males courted not only the model females of the same type, but also those of the different type. The results suggest that (1) sexual isolation exists between these types of E. hecabe on Okinawa-jima, although it is not perfect; (2) this isolation is mediated by female recognition but not male recognition; and (3) the sexual isolation system between two types is asymmetrical, depending on the female age after adult eclosion.     

Male mate-locating behavior inEuphydryas chalcedona (Lepidoptera: Nymphalidae) related to pupation site preferences

Males of the checkerspot butterfly (Euphydryas chalcedona)patrol and perch near but not on the larval foodplant in search of females. Experiments with tethered butterflies show that searching males chase virgin females for longer times than they do mated females or males. The larvae leave the larval food-plant to pupate. The correspondence between the distance from the larval foodplant to pupation sites and where males search for females suggests that male mate-locating behavior has evolved to maximize the rate of encounters with newly emerged, virgin females. These conclusions are compared to a recent analysis of butterfly mating systems by Odendaal et al. (Am. Nat.125: 673–678, 1985).     

Brood Odor Discrimination Abilities in Hygienic Honey Bees (Apis mellifera L.) Using Proboscis Extension Reflex Conditioning

To understand the effect of abnormal brood odors on the initiation or control of hygienic behavior in honey bees, we employed the associative learning paradigm, proboscis extension reflex conditioning. Bees from two genetic lines(hygienic and non-hygienic) were able to discriminate between high concentrations of two floral odors equally well. Differential discrimination abilities were observed between the two lines when healthy and diseased brood odors were used, with the bees from the hygienic line discriminating between the pair of brood odors better than the non-hygienic bees. These results suggest that hygienic behavior in individual bees is associated with the bees' responses to olfactory stimuli emanating from diseased brood.     

Structure, mechanism and mechanical properties of pupal attachment in Greta oto (Lepidoptera: Nymphalidae: Ithomiinae)

The structure and mechanism of pupal attachment are described for the nymphalid Greta oto using electron microscopy, and high‐speed and time‐lapse photography. The cremaster is composed of a 3‐D array of hooked setae that engage with silk fibers spun into layers in a pad on the lower leaf surface. Each seta comprises a shaft terminating in a strongly curved hook, tipped with two lateral barbs. These hook into the silk pad, which is densely laid and built‐up in the central portion, flattening out peripherally. Time‐lapse photography showed that silk pad construction by fifth instar larvae is completed in four distinct spinning movements, producing a random fiber arrangement. It is proposed that such a fiber arrangement provides isotropic strength, giving greater flexibility to the attachment. The cremaster is attached to the silk pad by a series of lateral movements of the pupa's posterior abdomen. This movement, together with the shape of the setal hooks, is thought to be integral to the attachment process. Tensile loading tests showed that attachment failure is due to the breakage of the silk pad, which undergoes gradual destruction before releasing the cremaster. The attachment was found to have high tensile strength and fracture toughness, both of which suggest that it has evolved for the dual purpose of preventing the pupa being pulled from the leaf by a predator and preventing the attachment being weakened by wind, which causes the pupa to swing.     

Investigating the Consistency of Mate-Locating Behavior in the Territorial Butterfly Hypolimnas bolina (Lepidoptera: Nymphalidae)

The study of butterfly behavior has afforded valuable insights into the evolution of alternative mating tactics. Two hypotheses derived from this area of research contend that (1) territoriality is only viable under low to moderate conspecific densities (due to the costs of site defence) and (2) perching may be employed only when thermal conditions constrain flight activity. These hypotheses were evaluated by investigating mate locating behavior in Hypolimnas bolina, a territorial species that is naturally subject to variation in population density and weather conditions. Male behavior was charted throughout the day during a period of high population density at an encounter site in tropical Australia. Perching was the primary tactic, although a small proportion of individuals patrolled nonaggressively in the afternoon. Population-level male behavior failed to support predictions drawn from either the territory economics or thermal constraint hypotheses. First, the proportion of perching males and the number of aggressive conspecific interactions (per male) increased with increasing male density at the site. Second, few males patrolled at the hottest, brightest time of day (approximately midday), and the diel distribution of perchers did not emulate the U-shaped distribution shown by the occurrence of dorsal basking behavior. These results show that perching in this species is not a suboptimal tactic employed when temperatures constrain flight activity but may represent the best method of locating receptive females. At this stage the reproductive significance of the observed patrolling behavior remains obscure.     

The Functional Role of the Hollow Region of the Butterfly Pyrameis atalanta （L.） Scale

Questions concerning the functional role of the hollow region of the butterfly Pyrameis atalanta （L.） scale are experimentally investigated. Attention was initially directed to this problem by observation of the complex microstrucmre of the butterfly scale as well as other studies indicating higher lift on butterfly wings covered with scale. The aerodynamic forces were measured for two oscillating scale models. Results indicated that the air cavity of an oscillating model of the Pyrameis atalanta （L.） scale increased the lift by a factor of 1.15 and reduced the damping coefficients by a factor of 1.38. The modification of the aerodynamic effects on the model of butterfly scale was due to an increase of the virtual air mass, which influenced the body. The hollow region of the scale increased the virtual air mass by a factor of 1.2. The virtual mass of the butterfly scale with the hollow region was represented as the sum of air mass of two imaginary geometrical figures： a circular cylinder around the scale and a right-angled parallelepiped within the hollow region. The interaction mechanism of the butterfly Pyrameis atalanta （L.） scale with a flow was described. This novel interaction mechanism explained most geometrical features of the airpermeable butterfly scale （inverted V-profile of the ridges, nozzle of the tip edge, hollow region, and openings of the upper lamina） and their arrangement.     

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.     

Olfactory responses of Aphidius gifuensis to odors of host plants and aphid‐plant complexes

Abstract The olfactory responses of Aphidius gifuensis to odors from two host plants (Nicotiana tabacum and Brassica napus ssp.) and their complexes with different infestation levels of two host aphids (Myzus persicae and Lipaphis erysimi) were respectively examined in an olfactometer. The results showed that female A. gifuensis did not respond to odors of undamaged or mechanically damaged host plants, but significantly responded to odors of aphid/plant complexes. Moreover, A. gifuensis responded significantly to odors of both M. persicae and L. erysimi/plant complexes when host plants were infested by high levels of aphids, suggesting that quantity of aphid‐induced volatiles could be important for attracting A. gifuensis. When tested between aphid/plant complexes, A. gifuensis did not show its preference for either complex. The efficiency of A. gifuensis against aphids in open fields potentially could be improved by using its olfactory response to aphid/plant complexes.