Pupal colour dimorphism in California Battus philenor: pupation sites, environmental control, and diapause linkage

ABSTRACT.

• 1 Natural pupation sites and corresponding pupal colour (green or brown) were determined for samples of Battus philenor (L.) from two Californian populations.
• 2 Larvae pupate off the ground on trees, shrubs and man-made objects.
• 3 The vertical distribution of pupation sites and relative frequencies of pupae formed on narrow twigs and broad substrates show interpopulation variability, and seem to be determined by habitat-specific and possibly behavioural differences among populations.
• 4 The percentage of‘mismatched’pupae in green leafy environments (brown) is greater than that on wide substrates (green). Heterogeneity in samples of the latter suggest strong but sporadic predation pressure on non-cryptic pupae in exposed areas.
• 5 Green and brown substrates generally promoted formation of cryptic green and brown pupae although rearing conditions modified pupal colour response to substrate colour and larval pupation site choice.
• 6 Warm temperatures and long days increased the production of brown pupae. Short photoperiods increased the tendency of larvae to pupate on narrow twig-like substrates and to form green pupae.
• 7 Green pupae show less tendency to diapause than brown pupae. The difference between percentage diapause in the two colour forms increases under conditions favouring progressively more continuous development.

     

Regulatory mechanisms in phenotypic plasticity of diapause and nondiapause pupal colouration of the swallowtail butterfly Papilio machaon

Nondiapause pupae of Papilio machaon L. exhibit pupal colour diphenism comprising green–yellow and brown–white types. To understand the regulatory mechanism underlying the control of pupal colouration in P. machaon, the effect of environmental cues on diapause and nondiapause pupal colouration is investigated. When larvae reared under short‐day and long‐day conditions are allowed to pupate in sites with a smooth surface and a yellow background colour, all diapause pupae exhibit a brown–white type and 89.5% of nondiapause pupae exhibit a green–yellow type, respectively. With rough‐surface pupation sites, all diapause pupae exhibit brown–white and intermediate types, whereas a large proportion of nondiapause pupae exhibit brown–white and intermediate types, although some exhibit a green–yellow type. When extracts prepared from the head‐thoracic and thoracic‐abdominal regions of larval central nervous systems are injected into the ligated abdomens of P. machaon short‐day pharate pupae, all recipients exhibit a brown–white colouration. Furthermore, when each extract is injected into the ligated abdomen of Papilio xuthus L. short‐day pharate pupae with orange‐pupa‐inducing factor activity, recipients injected with the head‐thoracic extract exhibit the brown type, whereas those injected with the thoracic‐abdominal extract exhibit an orange colour. The results indicate that the response to the environmental cues of pupation site in P. machaon changes according to the photoperiodic conditions experienced during larval stages, and that at least two hormonal factors producing brown–white pupae are located in the larval central nervous system, with the secretion of these factors being regulated by the recognition of environmental cues in long‐day larvae.     

Pupal colour dimorphism in a desert swallowtail (Lepidoptera: Papilionidae) is driven by changes in food availability,not photoperiod

1. The swallowtail butterfly Battus polydamas archidamas Boisduval, 1936, exhibits polyphenism for pupal coloration (green and brown). It is distributed across arid regions with winter rains and is monophagous on Aristolochia plants, which emerge after the winter rains and dry out the during summer. Thus, day length does not covary positively with host plant productivity. It was hypothesised that pupal colour was driven by food availability, not photoperiod. The benefits of pupal coloration matching the colour of pupation sites in terms of field survival were also investigated to evaluate the adaptive value of pupa colour. 2. Larvae were reared under a factorial array of two photoperiods (LD 10:14 h and LD 14:10 h) and two food availability regimes (leaves ad libitum and available every other day) to assess the frequency of green and brown pupae. Field survival of green and brown pupae was quantified in three commonly used habitats that differ in background coloration (cacti, rocks and shrubs). 3. Food availability determined pupal colour. Larvae in the ad libitum regime resulted mostly in green pupae, while those with restricted food were mostly brown. In contrast, photoperiod did not influence pupal colour. Survival probability of pupae placed on cacti was higher than those placed on rocks and shrubs, and the lowest predation risk across habitats was for green pupae on cacti. 4. Food availability plays a major role in the seasonal polyphenism for pupal colour of specialist butterflies inhabiting arid environments with winter rains.     

Foodplant effects on colour morphs of Eumorpha fasciata caterpillars (Lepidoptera: Sphingidae)

Caterpillars of the hawkmoth Eumorpha fasciata are highly polymorphic for colour, with green, pink, and pink-and-yellow forms in the second through fourth instars, and green and multicoloured forms in the fifth instar. Four years of field censuses on four foodplant species determined that all morphs were found on all plant species; morph frequencies were homogeneous on each plant species over time; and morph frequencies differed consistently among plant species. When larvae were reared from eggs on three of the hostplant species in the laboratory, differences in morph frequencies paralleled the census results. Thus foodplant quality is one factor affecting larval colour in E. fasciata. A literature survey reveals that foodplant effects on larval colouration may be widespread in the family Sphingidae, but most reports are anecdotal rather than experimental. The implications of this mechanism of colour determination are discussed.     

Factors affecting pupal survival and eclosion in the pine beauty moth,Panolis flammea (D&S)

Summary The proportion of adult Panolis flammea emerging from the overwintering pupae was markedly affected by pupation substrate, waterlogging, temperature and relative humidty. Pupae which had spent the winter in needle litter had a significantly greater survival rate than those in either soil or peat.The greater the length of time spent waterlogged the greater the mortality rate of P. flammea pupae. At temperatures of-20° C, there was 100% pupal mortality within 24 h of exposure.Pupal weight loss was proportional to the length of time spent as a pupa. Female pupae lost proportionately more weight than male pupae. Pupae of both sexes lost more weight at a low relative humidity than at a high relative humidity. At low relative humidities female pupal mortality was higher than that of male pupae.     

Gregarious pupation act as a defensive mechanism against cannibalism and intraguild predation

Coccinellid pupae use an array of defensive strategies against their natural enemies. This study aims to assess the efficiency of gregarious pupation as a defensive mechanism against intraguild predators and cannibals in coccinellid. The study was designed specifically (i) to determine the natural occurrence of gregarious pupation in the field for different coccinellid species, and (ii) to evaluate the adaptive value of gregarious pupation as a defensive mechanism against 2 types of predators (i.e., cannibals and intraguild predators). In the field, gregarious pupation consisted of a group of 2–5 pupae. The proportion of gregarious pupation observed varied according to species, the highest rate being observed with Harmonia axyridis Pallas (Coccinellidae; 14.17%). Gregarious pupation had no impact on the probability that intraguild predators and cannibals locate pupae. Intraguild predation occurred more often in site with gregarious pupation, while cannibalism occurred as often in site with gregarious pupation as in site with isolated pupa. However, for a specific pupa, the mortality rate was higher for isolated pupae than for pupae located in a gregarious pupation site both in the presence of intraguild predators and in the presence of cannibals. The spatial location of pupae within the group had no impact on mortality rate. Since it reduces the risk of predation, it is proposed that gregarious pupation act as a defensive mechanism for H. axyridis pupae.     

Concurrently burrowing host fly larvae reciprocally enhance pupation depth to minimise parasitism risk

1. Variation in parasitism risk among hosts is a key factor influencing host–parasitoid interactions; however, within-patch variation (as opposed to between-patch variation) in parasitism risk has hardly been studied. This study investigated the mechanisms of within-patch variation in parasitism risk in the oriental fruit fly, Bactrocera dorsalis (Hendel) (Diptera: Tephritidae). 2. Bactrocera dorsalis pupates underground. Previous studies have shown that pupae at greater depths have a reduced risk of parasitism by a pupal parasitoid Dirhinus giffardii (Silvestri) (Hymenoptera: Chalcididae). When a pupa near the ground surface is parasitised, the victim may decrease the risk of parasitism risk for pupae located at deeper depths. It was hypothesised that larvae will pupate at greater depths when they perceive the presence of conspecifics because of the benefit of pupating deeper than other individuals. 3. In a series of laboratory experiments, the effects of three factors on pupation depth were examined, including: (i) the density of larvae during development; (ii) the density of concurrently burrowing larvae; and (iii) the presence of pre-existing pupae in the pupation substrate. Only the density of concurrently burrowing larvae influenced pupation depth, which suggests that when a burrowing larva perceives the presence of other burrowing larvae, it aims to burrow deeper than them. 4. This study shows not only that parasitism risk is variable among pupae within a patch, but also that it is density-dependent. A commonly made assumption (i.e. the absence of within-patch variability in parasitism risk) needs to be re-evaluated in a wide range of host–parasitoid systems.     

Convergent evolution of neuroendocrine control of phenotypic plasticity in pupal colour in butterflies

Phenotypic plasticity in pupal colour occurs in three families of butterflies (the Nymphalidae, Papilionidae and Pieridae), typically in species whose pupation sites vary unpredictably in colour. In all species studied to date, larvae ready for pupation respond to environmental cues associated with the colour of their pupation sites and moult into cryptic light (yellow–green) or dark (brown–black) pupae. In nymphalids and pierids, pupal colour is controlled by a neuroendocrine factor, pupal melanization-reducing factor (PMRF), the release of which inhibits the melanization of the pupal cuticle resulting in light pupae. In contrast, the neuroendocrine factor controlling pupal colour in papilionid butterflies results in the production of brown pupae. PMRF was extracted from the ventral nerve chains of the peacock butterfly Inachis io (Nymphalidae) and black swallowtail butterfly Papilio polyxenes (Papilionidae). When injected into pre-pupae, the extracts resulted in yellow pupae in I. io but brown pupae in P. polyxenes. These results suggest that the same neuroendocrine factor controls the plasticity in pupal colour, but that plasticity in pupal colour in these species has evolved independently (convergently).     

Natural pupation sites of swallowtail butterflies (Lepidoptera: Papilioninae): Papilio polyxenes Fabr., P.glaucus L. and Battus philenor (L.)

Abstract. 1. Natural pupation sites have been found in Papilio polyxenes and P.glaucus by releasing prepupal larvae marked with UV-fluorescent paint and locating them at night with a UV lamp, and in Battus philenor by searching a forest habitat where the larval foodplant is abundant.
2. P.polyxenes , a species of weedy habitats, pupates off the ground on a variety of substrates including grasses, weed stalks, posts, etc. The pupae may be green or brown, resembling the substrate.
3. P.glaucus , a species of forest habitats, pupates very close to the ground in the litter and has monomorphic brown pupae.
4. B.philenor , also a forest species, pupates on exposed surfaces (chiefly tree-trunks or cliffs) well off the ground. Its pupae may be brown or green, but the latter were found only on the slenderest twigs.
5. The results for polyxenes and glaucus support the generalization of Clarke & Sheppard (1972) that species of stable habitats are likely to have monomorphic pupae, while those of habitats in which available sites may not be so similar from one generation to the next will be dimorphic.
6. B.philenor is more problematical, but its tendency towards pupal monomorphism (brown) is logical in relation to its common pupation sites.     

<Emphasis Type="Italic">Drosophila</Emphasis> pupation behavior in the wild

We investigated pupa distributions of D. simulans, D. buzzatii, D. melanogaster, D. immigrans and D. hydei on a number of natural breeding sites. Pupae of all five species showed aggregated distributions, which prompted us to examine these aggregations in a more detail for two species that commonly co-occur in breeding sites, D. simulans and D. buzzatii. We found that pupae of both species tend to be aggregated in conspecific clusters. Subsequent experiments revealed that both species are attracted to the odors of other larvae, though only D. buzzatii differentiated between conspecifics and heterospecifics (they preferred conspecific). Furthermore, third instar larvae of both species preferred more alkaline substrates. Altogether, our results demonstrate that Drosophila species form conspecific pupa aggregations in natural breeding sites, and that pupation site selection depends on interactions among conspecific and heterospecific larvae and on chemical characteristics of the breeding sites.     

Dispersal and Burial Behavior in Larvae of <Emphasis Type="Italic">Chrysomya megacephala</Emphasis> and <Emphasis Type="Italic">Chrysomya albiceps</Emphasis> (Diptera,Calliphoridae)

Blowflies use discrete and ephemeral substrates to feed their larva. After they run out of food, the larvae begin to disperse in order to find adequate places for pupation or additional food sources, a process named post-feeding larval dispersal. Briefly state the aspects and why they are important were studied in a circular arena of 25 cm in diameter and covered with wood shavings to a height of 40 cm allowing post-feeding dispersal from the center of the arena. Larvae of both Chrysomya albiceps and C. megacephala were used in five experiments for each species. For each pupa location, determined as distance from the center, depth, and weight were evaluated. Statistical tests were done to verify the relation between weight, depth and distance for pupation and for larvae of two species shows that the media distance is significantly different for two species and for C. megacephala this distance is greater than the distance for C. albiceps. The depth too is different for each species, as the larvae of C. megacephala buries deeper than C. albiceps. With relation of weight, there is no statistic evidence that have any difference between weights for pupation for each species.     

New data on the ecology and morphology of xylobiont larvae of the genus <Emphasis Type="Italic">Elephantomyia</Emphasis> Ost.-Sack. (Diptera,Limoniidae)

New data on the biology and morphology of the larva and pupa of E. hokkaidensis Al. and E. krivosheinae Sav. are given. Additional data on E. edwardsi Lack. and E. subterminalis Al. distributed in Russia are given for the first time. Keys to the larvae and pupae are compiled.     

The evolution of environmentally-cued pupal colour in swallowtail butterflies: natural selection for pupation site and pupal colour

1. Environmentally-cued pupal colour in swallowtail butterflies has been hypothesized to evolve as a consequence of (a) the evolution of a preference for pupation sites above the ground that vary in colour and (b) natural selection for crypsis on such sites.
2. This hypothesis was tested by comparing the field survival of green and brown Papilio polyxenes Fabr. pupae placed on green or brown pupation sites that were either above the ground on near the ground.
3. Green pupae on green sites above the ground had a significantly higher probability of survival than did all other pupal colour and pupation site combinations.
4. Pupae on sites above the ground were more likely to be preyed upon during the day, whereas those on sites near the ground were more likely to be preyed upon during the night, suggesting that variation in nocturnal and diurnal predation influences the evolution of pupation site preference.
5. To the extent that diurnal predators use colour vision to locate prey, diurnal predation should favour environmentally-cued pupal colour.     

Action spectra for light-controlled pupal pigmentation in Pieris brassicae: Melanization and level of bile pigment

Pupal pigmentation in Pieris brassicae is controlled by light during a sensitive period before pupation. Action spectra for melanization and level of biliverdin-IX_λ were determined in the range of 383 to 765 nm at equal quantum flux densities. Darkness and light of 500, 728, and 765 nm result in intermediate melanization and a low level of bile pigment. Wavelengths of <500 nm promote, while those of >500 nm inhibit cuticular melanization, but the level of bile pigment is raised in both. Infrared light (748 nm) is shown to have a significant effect, but only the bile pigment is influenced. In both spectra, maximal effects are observed at identical wavelengths: 404, 577, and 661 nm; a minimum occurs at 640 nm. The degree of melanization and the level of biliverdin are correlated positively for λ<500 nm and negatively for λ>500 nm. It is argued that the action spectra may not reflect the absorption of the photoreceptor(s), but rather the action of mechanisms in the central nervous system, which influence both pigments in separate and different ways. Light does not stimulate the synthesis of bile pigment in the larva before pupation, but does inhibit its degradation in the pupa. The results are discussed in the light of the endocrine mechanisms involved, and in respect of the biological significance of colour adaptation in Pieris pupae.     

The use of visual cues in host evaluation by aphidiid wasps I. Comparison between threeAphidius parasitoids of the pea aphid

Host evaluation behaviour was examined in three species of aphid parasitoids,Aphidius ervi haliday,A. pisivorus Smith, andA. smithi Sharma & Subba Rao (Hymenoptera: Aphidiidae). Parasitoids were provided under laboratory conditions with three kinds of hosts representing two aphid species: (green) pea aphid,Acyrthosiphon pisum (Harris), and green and pink colour morphs of the alfalfa aphid,Macrosiphum creelii Davis. Females of all threeAphidius species distinguished between aphids on the basis of colour, movement, and host species. Patterns of host acceptance by parasitoids were species-specific. InA. ervi, host preference was the same in light and dark conditions: pea aphid>green alfalfa aphid≫pink alfalfa aphid. In contrast,A. pisivorus attacked and accepted pea aphid and green alfalfa aphid equally in the light and preferred both of these over pink alfalfa aphid; however, it made no distinction between pea aphid and pink alfalfa aphid in the dark. Females ofA. smithi attacked all three kinds of hosts (pea aphid>green alfalfa aphid≫pink alfalfa aphid) but apparently laid eggs only in pea aphid. The frequencies of attack and oviposition by all wasps were higher on ‘normal’ pea aphids than on those anaesthetized with CO₂. Host recognition is confirmed by chemical cues in the aphid cuticle that are detected during antennation, and host acceptance is dependent on an assessment of host quality during ovipositor probing.     

Effects of short-term heat stress on the growth and development of Bradysia cellarum and Bradysia impatiens

Bradysia cellarum Frey and Bradysia impatiens Johansen are major pests of vegetable crops, as well as edible mushrooms and ornamental plants, and damage to hosts resulting in economic losses. Temperatures above the optimum levels for these pests have been predicted to regulate their population growth during summer. The aim of the present study was to examine the effects of both heat stress and exposure time on the growth and development of eggs, larvae and pupae for two Bradysia species. The egg stage, egg hatching rate, 4th instar larval stage, pupation rate, pupal stage and adult emergence rate were observed after exposing at high temperatures of 34°C, 37°C and 40°C for 1, 2, 4 and 6 hr. The results showed that 34°C, 37°C and 40°C for 1-, 2-, 4- and 6-hr exposure treatments prolonged the developmental stage of egg, 4th instar larva and pupa, while decreasing the egg hatching rate, pupation rate and adult emergence rate. This suggests that increasing temperature or prolonging exposure time to the heat stress could significantly affect insect survival, growth and development. Our study could provide an ecological basis for pests’ management strategy by using short-term heat stress.     

The Pupa and Larva of Propsilocerus taihuensis (Wen,Zhou & Rong) (Diptera: Chironomidae), with Complete Keys to all Immatures of the Genus Propsilocerus

The pupa and larva of Propsilocerus taihuensis (Wen, Zhou & Rong, 1994) are described for the first time. Both are close to P. akamusi (Tokunaga). The phylogenetic argumentation of the genus outlined in Sæther and Wang (1996) is confirmed. Revised keys to larvae and pupae of the genus are given. The main morphological characters of the different species of the genus are compared in tables.     

Scirtes hemisphaericus uses macrophyte snorkels to pupate under water. With notes on pupae of additional European genera of Scirtidae (Coleoptera)

Morphological and behavioural adaptations enable Scirtes hemisphaericus to pupate under water, using emergent macrophytes as snorkels to breathe atmospheric air. The last instar larva bites into emergent macrophytes until the aerenchym is exposed. The pupa pushes its fore body into the wound, becoming surrounded by an air film in continuity with intracellular and atmospheric air. The pupa lacks normal ecdysial sutures, instead, its cuticle ruptures between the pro- and mesothorax. The fore body cuticle remains in place, preserving the connection to atmospheric air for the immature beetle. Pupation in earthen cells is known only in Elodes. Several other genera pupate under some cover, or freely exposed. Most pupae have 2–4 horn-like pronotal processes and 2–4 caudal attachment organs which differ between genera and sexes. Hydrocyphon (which pupates under water) and S. hemisphaericus pupae lack pronotal horns and attachment organs.