Can life‐history and defence traits predict the population dynamics and natural enemy responses of insect herbivores?

Abstract.  1. Life-history differences between herbivorous insects with eruptive and latent population dynamics are potentially useful for predicting population size variability. An association has also been demonstrated between herbivorous insect defence traits and the responses of various natural enemies.
2. Here predictions of population dynamics and natural enemy responses based on life-history and defence traits are tested using Gonometa postica Walker and G. rufobrunnea Aurivillius, two Southern Hemisphere Macrolepidoptera (Lasiocampidae) species. The temporal and spatial variation in pupal abundance and patterns of pupal parasitism and predation for both species are described and quantified for the first time.
3. Eleven sites were sampled over four generations across the region where both species have historically reached high population densities. Although there was evidence suggesting that population synchrony is driven by weather patterns, site-specific environmental differences contributed to the observed population variability. This study is the first to quantify the extent of population size variability of a species with an intermediate position on the eruptive–latent population dynamic gradient, where data on insect population dynamics is scarce.
4. Support for the life-history–population dynamic relationship was found, as intermediate population size variability for these species was observed. Larval and pupal defence traits, however, were poor and inconsistent predictors of mortality rate. Pupal cocoon structure differences, previously documented for these Gonometa species, may in fact explain the interspecific differences in natural enemy responses found.
5. Predicted population dynamics and natural enemy responses may, however, be overridden by ecological conditions. Nevertheless, life-history and defence traits provide a useful basis for predicting population dynamics of poorly studied species.     

Two different sensory mechanisms for the control of pupal protective coloration in butterflies

The butterflies Graphium sarpedon nipponum Fruhstorfer and Papilio xuthus Linné show pupal protective color polymorphism, but the two species appear to have different sensory mechanisms for determining pupal coloration. When light was of sufficient illumination, the larvae of Graphium sarpedon became bright yellowish green pupae on white pupation boards and reddish brown pupae on black pupation boards. The pupal coloration thus strongly depended on the brightness of the pupation site. In addition, larvae became bright yellowish green pupae in complete darkness. From these results, measurement of the illumination suggested that pupal color is determined by the illuminant difference between incidence light from the dorsal direction and ventral light from a paper board; i.e., the sum of the reflected light of the board plus the penetrated light passing through the board. The illuminant difference required for reddish brown coloration was 40 lux or more. The optical signals received through the stemmata during a critical period before formation of the thorax garter (band string) were important for coloration. By contrast, in Papilio xuthus, successive tactile signals from a rough surfaced pupation site during a critical period before and after formation of the garter were important for determining brown pupal coloration.     

Interactions of environmental factors influencing pupal coloration in swallowtail butterfly Papilio xuthus

The swallowtail butterfly Papilio xuthus Linné [Lepidoptera: Papilionidae] exhibits pupal protective color polyphenism. Interactions of various environmental factors on pupal coloration were analyzed in non-diapausing individuals. Under sufficient light (200lux), most pupating larvae became green pupae when the surface of the pupation site was smooth, while they became brown when the surface was rough. Tactile signals are the positive environmental factors causing induction of the brown pupal coloration. In dark boxes, the induction of the brown pupal coloration was easily induced even on a smooth surface, suggesting that light suppresses induction of brown coloration. Different colors of pupation sites did not affect pupal coloration under sufficient light. Environmental factors received during a critical period both before girdling and after girdling affected pupal coloration. When tactile signals received from rough surfaces reach threshold levels during pupation, brown pupal coloration is determined. Larvae reared under a daily periodicity of natural light formed a girdle at midnight, subsequently, the prepupae received strong daylight the following day. Under natural light most larvae produced brown pupae on rough surfaces and green pupae on smooth surfaces.     

Closely related parasitoids induce different pupation and foraging responses in Drosophila larvae

Few examples exist where parasites manipulate host behaviour not to increase their transmission rate, but their own survival. Here we investigate fitness effects of parasitism by Asobara species in relation to the pupation behaviour of the host, Drosophila melanogaster . We found that Asobara citri parasitized larvae pupate higher in rearing jars compared to unparasitized controls, while A. tabida pupated on or near the medium. No change in pupation site was found for three other species. A follow-up experiment showed a non-random distribution of parasitized and unparasitized pupae over the different jar parts. To test the adaptiveness of these findings, we performed pupal transfer experiments. Optimum pupation sites were found to be different between host individuals; wall individuals survived better than bottom individuals, but bottom individuals did worse at the wall. Two parasitoid species that alter pupation site significantly showed high rates of diapause at their 'preferred' pupation site. For one of them, A. citri , pupation occurred at the optimal site for highest survival (emergence plus diapause). From literature we know that pupation height and foraging activity are genetically positively linked. Therefore, we implement a short assay for rover/sitter behavioural expression by measuring distance travelled during foraging after parasitism. For one out of three species, foraging activity was reduced, suggesting that this species suppresses gene expression in the for pathway and thereby reduces pupation height. The parasitoid species used here, naturally inhabit widely different environments and our results are partly consistent with a role for ecology in shaping the direction of parasite-induced changes to host pupation behaviour. More parasitoids are found on the wall of the rearing jar when they originate from dry climates, while parasitoids from wet climates pupate on the humid bottom.     

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).     

Pupation Site Selection in Four Drosophilid Species: Aggregation and Contact

This study of pupation site selection was undertaken to assess the frequency and orientation of pupae making physical contact in four sympatric species in the family Drosophilidae: Drosophila melanogaster, D. simulans, D. funebris, and Zaprionus tuberculatus. Pupation behavior was assayed in a vial containing a small food cup. Using nearest-neighbor analysis, we found that pupae were aggregated. Furthermore, the frequency of contact between pupae was far greater than could be explained by chance, in all four species. In the three species of Drosophila, about a third of the contacts were between intimately paired pupae whose long axes were parallel; we call this arrangement synapsis. In Z. tuberculatus, most pupal contacts were unoriented. When D. melanogaster larvae were reared with each of the other species, heterospecific pupal contact, including synapsis, occurred. Our discovery of pupal contact in several drosophilids expands the known repertory of social behavior in this family.     

Incorporating biotic interactions in the distribution models of African wild silk moths (Gonometa species,Lasiocampidae) using different representations of modelled host tree distributions

Biotic interactions influence species niches and may thus shape distributions. Nevertheless, species distribution modelling has traditionally relied exclusively on environmental factors to predict species distributions, while biotic interactions have only seldom been incorporated into models. This study tested the ability of incorporating biotic interactions, in the form of host plant distributions, to increase model performance for two host‐dependent lepidopterans of economic interest, namely the African silk moth species, Gonometa postica and Gonometa rufobrunnea (Lasiocampidae). Both species are dependent on a small number of host tree species for the completion of their life cycle. We thus expected the host plant distribution to be an important predictor of Gonometa distributions. Model performance of a species distribution model trained only on abiotic predictors was compared to four species distribution models that additionally incorporated biotic interactions in the form of four different representations of host plant distributions as predictors. We found that incorporating the moth–host plant interactions improved G. rufobrunnea model performance for all representations of host plant distribution, while for G. postica model performance only improved for one representation of host plant distribution. The best performing representation of host plant distribution differed for the two Gonometa species. While these results suggest that incorporating biotic interactions into species distribution models can improve model performance, there is inconsistency in which representation of the host tree distribution best improves predictions. Therefore, the ability of biotic interactions to improve species distribution models may be context‐specific, even for species which have obligatory interactions with other organisms.     

Biology of Xylobiont larvae of limoniid flies of the genus Gnophomyia (Diptera, Limoniidae) with description of immature stages

New data on the larval and pupal morphology and the life history of limoniid flies (G. lugubris (Zett.), G. acheron Al., and G. viridipennis (Gimmerthal)) of the genus Gnophomyia are presented. Larvae of all the species are phloeophages and inhabit tree bark. Keys to larvae and pupae of these species are given.     

Environmental control of pupal colour in swallowtail butterflies (Lepidoptera: Papilioninae): Battus philenor (L.) and Papilio polyxenes Fabr.

Abstract. 1. Some swallowtail butterflies produce both green and brown pupae. The phenotypes result from the joint action of genotype and environment and usually make the pupae cryptic in their habitats.
2. The major environmental cues influencing pupal colour in two swallowtail species were determined to be textural and optical.
3. Differences in the usage of these kinds of cues in the two species are thought to have evolved because of major differences in the pupation habitats. P.polyxenes , which usually pupates on slender stems amidst vegetation, responds more strongly to optical cues. B.philenor , which usually pupates on exposed surfaces of tree trunks and cliffs, responds more strongly to textural cues.
4. Differences in the overall tendency to produce brown pupae ('sensitivity': Hazel, 1977) are thought to be related to the frequency of brown pupation sites utilized by these two species: high average sensitivity in philenor , which often uses brown sites, and lower average sensitivity in polyxenes , which often uses green sites.     

Population biology of the broad-horned flour beetle,Gnathocerus cornutus (F.) II. Crowding effects of larvae on their survival and development

Crowding effects of larvae on survival and development were examined for the broad-horned flour beetle, Gnathocerus cornutus (F.). The larvae matured about 3 weeks after hatching regardless of their densities, but pupation was severely hindered by crowding. There existed an upper limit for the number of the pupae produced and its mechanism was studied by a statistical analysis of the distribution patterns of pupal cells and the experiment in which glass tubes were artificially supplied in addition as pupation site. These studies show that G. cornutus larvae have a habit to construct cells for pupation and this habit leads to a contest competition for pupation site at high densities. The significance of the contest competition for population regulation was discussed comparing the results on Tribolium confusumJacqueline duVal .     

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.     

Comparing the rheology of mulberry and "wild" silkworm spinning dopes

Lepidoperan silks provide a superb opportunity for comparative studies of spinning and fiber characteristics. Comparing the four species, Bombyx mori (China), Actias selene (India), Antheraea yamamai (Japan), Gonometa postica (Africa), allows us to examine differences on the family, species, and race levels. Measured rheological properties were consistent with phylogenetic relationships and in the context of resource allocation and gland morphology. We propose that the thorough domestication of the mulberry silkworm B. mori for high silk yield has resulted in a compensatory optimization for spinning efficiency. This is in stark contrast to the wild silkworms, where Saturnids appear to minimize their energetic input toward silk output and G. postica seems to balance both. We conclude that comparative studies provide valuable baseline information for future biomimetic applications and modeling, as well as illuminating biologically important details of silk processing.     

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 warning color development in above-ground pupating species but cryptic color in ground-surface pupating species of the nine-spotted diurnal moths (Erebidae: Arctiinae: Syntomini)

Insects usually have cryptic colors to avoid detection by visually hunting predators. However, if the insects acquire toxic or repellent substances against predators, some of them develop conspicuous coloration to exhibit their unpalatability. Such warning colors allow insects to survive. In the nine-spotted diurnal moths (Erebidae: Arctiinae: Syntomini), we found the above-ground pupating species to have conspicuous colored pupae, but the ground-surface pupating species to have cryptic colored pupae. In this study, the relationships between unpalatability and coloration of these pupae are examined among three species of Amata and one species of Syntomoides. Pupae of the two species (A. germana and A. flava) are conspicuous in their color pattern with seven black dotted lines longitudinally on their pale-yellow bodies. These pupae are exposed to the aerial predators in a coarse silk mesh hanging from leaves and/or branches. The other two species (A. fortunei and S. imaon) pupate in spaces under stones, fallen twigs and leaves on the ground surface, and the pupae in a coarse silk cocoon is cryptic dark brown. Their pupation site selections are reproduced in the rearing glass vessels. Palatability assessment using lizards as a potential predator suggests that pupae of A. germana, A. flava and A. fortunei are unpalatable and the lizard's feeding response decreases with experience. However, pupae of S. imaon are all eaten (palatable). Finally, the possible evolutionary scenario of pupal colors of these four species is discussed in relation to pupation site selection and palatability.     

Pupal polymorphism in the butterfly Danaus chrysippus (L.): environmental, seasonal and genetic influences

The pupae of the tropical butterfly Danaus chrysippus are either green or pink the switch being operated by a ‘greening’ hormone produced in the larval head. Both environmental and genetic cues are involved in controlling the endocrine mechanism. The environmental factors identified are of two distinct kinds: proximate factors influence pupal colour after the larva has selected its pupation site, whereas ultimate factors are effective at an earlier stage, either prompting choice of pupation site by the larva or priming pupation physiology in a particular direction. Genetic factors preadapt the larva to form a pupa which will be cryptic in the normal or average conditions, climatic or biogeographical, anticipated in its environment. The proximate factors demonstrated are background colour, darkness, light quality (wavelength) and humidity. There is some evidence that substrate texture may also be relevant. Ultimate factors are temperature, humidity and species of larval foodplant. Two closely linked gene loci which govern the phenotype of adult morphs and races either have a pleiotropic effect on pupa colour or are closely linked with other genes which do so. Moreover, the two loci interact epistatically with respect to their pupation effects. Factors producing predominantly green pupae are plant substrates, yellow background, darkness, yellow light, high humidity, high temperature, the b allele at the B locus when homozygous and, on non-plant substrates, the C allele at the C locus. High frequencies of pink pupae result on non-plant substrates, red backgrounds, in blue light, low humidity, low temperatures and in B- and cc genotypes. The C locus alleles, C and c, interact epistatically with the B alleles, their effect on choice of pupation site being determined by linkage phase. Of the two foodplants tested, Calotropis produced a high frequency of green pupae and Tylophora of pinks. The seasonal cycling of rainfall, temperature, availability or condition of foodplant, and gene frequencies are all correlated with oscillations in the frequencies of green and pink pupae. Though genotype influences pupa colour, all genotypes are capable of forming pupae of both colours. The variation can therefore be attributed to an environmental polyphenism superimposed upon a genetic polymorphism. The hormone producing green pupae emanates from the head during the prepupal period. Denied hormonal influence, the pupa is pink. Pupal colour is judged to be aposematic at close range and cryptic at distance.     

<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.     

Parasitoids of the African wild silkmoth,Gonometa postica (Lepidoptera: Lasiocampidae) in the Mwingi forests,Kenya

Gonometa postica Walker produces silk of high quality, but it is affected by parasitoids attack. A study on the parasitism of G. postica larvae and pupae on host and non‐host plants were undertaken for the first and second generations, corresponding to the long (March–May) and short (October–December) rainy seasons in 2006 at six field sites, three each in the Imba and Mumoni forests of Mwingi, eastern Kenya. All freshly spun cocoons of G. postica were sampled at each site from a total of 100 trees of host plants and other non‐host plants where they have migrated before pupation. The cocoons were kept individually in fine net‐sealed plastic vials to determine percentage parasitism. Two dipterans and four hymenopteran larval–pupal parasitoids were identified from the two forests. The most common parasitoids were Palexorista sp. (Diptera: Tachinidae) and Goryphus sp. (Hymenoptera: Ichneumonidae) with parasitism ranging from 1.8 to 32.7% and 2.2 to 7.5%, respectively. Parasitism varied significantly according to host or non‐host plants, seasons and sites. This study indicates that, of the six parasitoid species recovered, only two had a significant impact in reducing the quality of the cocoons.     

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.     

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.

     

Patterns of pupal mortality in field populations of Hydropsyche and Cheumatopsyche (Trichoptera: Hydropsychidae)

SUMMARY. 1. Up to 40% of hydropsychid pupal cases (from three stations on the Credit and Humber Rivers, Ontario, Canada) contained insects already dead when collected; chironomid infestation accounted for up to 82% of total mortality within a taxon from any one station. The remaining mortality appeared to be due to siltation.
2. For all taxa ( Cheumatopsyche Wallengren and four species of Hydropsyche Pictet), and at all stations, prepupae suffered significantly more chironomid-related mortality than did fully-developed pupae.
3. Chironomid infestation generally affected all species of Hydropsyche equally; at some stations, Cheumatopsyche pupae appeared to suffer less chironomid-related mortality than did co-existing Hydropsyche species.
4. Chironomid infestation affected a greater proportion of pupae at the station where the density of pupal cases (per sampling quadrat) was greatest.
5. Vertical distribution of the pupation site had no apparent influence on mortality attributed to either siltation or chironomid infestation.
6. Chironomid infestation varied seasonally; it was greatest in May and July-August at an upstream station, and peaked in June at the downstream stations.
7. Mortality attributed to siltation was relatively constant for all stages and taxa, at all stations, throughout the sampling programme.