Erato Phyllis Research Articles

First report on the characterization of the gut microbiota in Heliconius erato phyllis caterpillars (Lepidoptera-Nymphalidae) from southern Brazil.

First report on the characterization of the gut microbiota in Heliconius erato phyllis caterpillars (Lepidoptera-Nymphalidae) from southern Brazil.

Landscape ecology and urban spatial configuration: Exploring a methodological relationship. Application in Pelotas, Brazil

Urbanization has been expanding rapidly in Brazil, negatively impacting the natural environment. Achieving sustainable development requires understanding the interdependencies between the social and natural worlds and finding a balance between them. Resilient urban planning considering green infrastructure can generate socio-environmental benefits through ecological connections. To achieve this, an integrated study between the social and environmental spheres is required, which are often developed separately. Thus, this investigation aims to develop a socio-environmental analysis by adopting urban configuration and landscape ecology models and subsequently comparing their respective outcomes. To this end, two models of spatial networks were examined in a sector of Pelotas, Brazil, based on graph theory and centrality measures. The urban configuration model was used to analyze human mobility in urban networks, while the landscape ecology model was used to study the butterfly species Heliconius erato phyllis in ecological networks. Our findings show no significant correlation between the outcomes generated by the two models, indicating a lack of cohesion in the location and connection of green infrastructure in the study area. This methodology represents a valuable tool for urban planners and researchers seeking to gain new insights into the performance of urban green areas while considering both human and species-related factors.

Identification and Synthesis of a Macrolide as an Anti-aphrodisiac Pheromone from Males of Heliconius erato phyllis.

Investigation of the contents of the scent glands of the heliconiine butterfly Heliconius erato phyllis via gas chromatography/electroantennography revealed an unprecedented active compound. The males transfer this compound to females during mating. The structure of (2R,6E,10R)-2,6-dimethyl-6-undecen-10-olide, a derivative of geranylacetone, was proposed on the basis of infrared and mass spectrometry spectra and microderivatization and confirmed by racemic and stereoselective syntheses. Bioassays with the synthetic macrolide showed the repellency of this compound, termed phyllisolide, when applied to scent glands of females, identifying it as an anti-aphrodisiac pheromone.

Morphology and histology of vom Rath's organ in brush-footed butterflies (Lepidoptera: Nymphalidae).

Vom Rath’s organ, located at the distal end of the third segment of the labial palp, is one of the recognized synapomorphies of Lepidoptera (Insecta). Information about the structural and histological morphology of this organ is sparse. The structure of vom Rath’s organ in four species of Nymphalidae, three frugivorous: Fountainea ryphea (Charaxinae: Anaeini), Morpho helenor achillaena (Satyrinae: Morphini) and Hamadryas epinome (Biblidinae: Ageroniini), and the nectarivorous species Aeria olena (Danainae: Ithomiini) is described by means of scanning electron microscopy and histology. The species showed significant differences in the cavity shape, setal morphology and arrangement, opening shape and location, associated with the organization of cell groups, type of axon, and degree of development. These differences do not seem to be related to feeding habit. No cell groups were found in Actinote thalia (Heliconiinae: Acraeini) and Heliconius erato phyllis (Heliconiinae: Heliconiini), and for the first time the absence of vom Rath’s organ is documented in the clade Ditrysia. A terminology is proposed to improve understanding of the organ morphology, with an extensive analysis of the previous descriptions.

Antimicrobial resistance and genetic relationships of enterococci from siblings and non-siblings Heliconius erato phyllis caterpillars.

BackgroundStudies evaluating bacteria in insects can provide information about host–microorganism–environment interactions. The gut microbial community has a profound effect on different physiological functions of insects. Enterococcus spp. are part of the gut community in humans and other animals, as well as in insects. The presence and antimicrobial resistance profile of enterococci are well studied in different animals; however, data for Heliconius erato phyllis (Lepidoptera: Nymphalidae) do not yet exist. Therefore, the aims of this study were to evaluate the distribution of enterococcal species, their antimicrobial resistance profile and virulence genes, and the genetic relationships between enterococci isolated from fecal samples from sibling and non-sibling H. erato phyllis caterpillars collected from different sites in South Brazil.MethodsThree H. erato phyllis females were captured (two from a forest fragment and one from an urban area), and kept individually in open-air insectaries. Eggs were collected and caterpillars (siblings and non-siblings) were fed daily with Passiflora suberosa leaves. Fecal samples (n = 12) were collected from fifth-instar caterpillars, inoculated in selective medium, and 15 bacterial colonies were randomly selected from each sample. Enterococci were identified by PCR and MALDI-TOF, analyzed by disk diffusion antimicrobial susceptibility tests, and screened for resistance and virulence genes by PCR. The genetic relationships between the strains were determined using pulsed-field gel electrophoresis (PFGE).ResultsA total of 178 enterococci strains were identified: E. casseliflavus (74.15%; n = 132), E. mundtii (21.34%; n = 38), E. faecalis (1.12%; n = 2) and Enterococcus sp. (3.37%; n = 6). High rates of resistance to rifampicin (56%) and erythromycin (31%) were observed; 120 (67.41%) of the isolates showed resistance to at least one antibiotic and six (3.37%) were multidrug-resistant.None of the erythromycin-resistant strains was positive for the erm(B) and msrC genes. The virulence genes esp, ace, and gelE were observed in 35%, 7%, and 1% of the strains, respectively. PFGE separated the enterococci into 22 patterns, four being composed of strains from sibling caterpillars.ConclusionEnterococcus casseliflavus was the dominant species in fecal samples of fifth-instar caterpillars. Resistant enterococci strains may be related to environmental pollution or the resistome. The PFGE analysis showed genetic relationships between some strains, suggesting that the enterococci isolated from fecal samples of the sibling caterpillars might have come from common sources, e.g., via diet (herbivory) and/or vertical transmission (through the egg surface). Further studies will be conducted to better understand the role of Enterococcus in the microbial community of the gastrointestinal tract of these insects, and the mechanisms involved in acquisition and maintenance of enterococci.

Structures related to pheromone storage in alar androconia and the female abdominal scent gland of Heliconius erato phyllis, Heliconius ethilla narcaea, and Heliconius besckei (Lepidoptera: Nymphalidae: Heliconiinae).

We describe the morphology of alar androconia and the female abdominal scent gland of Heliconius erato phyllis, Heliconius ethilla narcaea, and Heliconius besckei. Androconial scales of Heliconius, which are arranged in overlapping wing bands, release pheromones during courtship, probably through vibratory movements of male wings over the female to induce her to mate. An antiaphrodisiac is produced by glands located in the valves of the male and is transferred during copulation to the yellow dorsal abdominal sac present in the virgin female, causing this sac to emit a scent that reduces the attractiveness of the female for courtship with other males. Stereomicroscopy, SEM, and TEM analyses were conducted to describe the morphology of the internal and external scales and the external abdominal scent sac. The findings revealed different sizes of external androconial scales and an internal group of porous structural vesicles that are probably related to the preservation of internal space, reception and storage of secretions, and elimination of volatiles when the male is actively involved in courtship. Translucent projections on the female abdominal scent sac create open reservoirs for the reception, storage, and emission of antiaphrodisiac volatiles along with stink clubs. Male valve denticles vary in form and probably attach securely to the female sac during mating, thus ensuring secretion transfer. These features are discussed in the context of a comparative analysis.

Plasticidad fenotípica en mandíbulas de Heliconius erato (Lepidoptera: Nymphalidae) inducida por diferentes plantas hospederas (Passifloraceae)

El objetivo fue analizar si el consumo de diferentes plantas influenciaba el tamaño/forma de las mandíbulas de larvas de lepidópteros oligófagas y si dicho consumo ocasionaba tasas de crecimiento diferenciadas entre el músculo aductor mandibular y las mandíbulas durante su ontogénesis. Para esto se usaron larvas de Heliconius erato criadas en cuatro plantas hospederas naturales, a las que se les tomaron medidas de longitud de cápsula cefálica, tamaño mandibular y área del músculo aductor. Para detectar eventuales diferencias de forma mandibular se empleó microscopía electrónica y morfometria geométrica, y para posibles cambios alométricos se realizaron comparaciones entre curvas de crecimiento mandíbula vs. músculo aductor de los individuos criados en las cuatro plantas hospederas. Se encontró que no existen diferencias de tamaño entre las mandíbulas ni en músculos aductores de individuos criados en las diferentes plantas pero la forma mandibular fue significativamente diferente, mostrando una relación directa con la planta hospedera usada como alimento. Generalmente, la mandíbula creció proporcionalmente más que el músculo aductor, cuyo crecimiento fue estable. El cambio más significativo fue el alargamiento de los dientes incisivos en mandíbulas de larvas criadas con Passiflora suberosa. Se sugiere que esto se debe a que las hojas de esta especie poseen mayor dureza comparadas con las otras y que el desarrollar dientes largos facilitaría su corte y consumo.

Subtle variation in size and shape of the whole forewing and the red band among co-mimics revealed by geometric morphometric analysis in Heliconius butterflies.

Heliconius are unpalatable butterflies that exhibit remarkable intra‐ and interspecific variation in wing color pattern, specifically warning coloration. Species that have converged on the same pattern are often clustered in Müllerian mimicry rings. Overall, wing color patterns are nearly identical among co‐mimics. However, fine‐scale differences exist, indicating that factors in addition to natural selection may underlie wing phenotype. Here, we investigate differences in shape and size of the forewing and the red band in the Heliconius postman mimicry ring (H. erato phyllis and the co‐mimics H. besckei, H. melpomene burchelli, and H. melpomene nanna) using a landmark‐based approach. If phenotypic evolution is driven entirely by predation pressure, we expect nonsignificant differences among co‐mimics in terms of wing shape. Also, a reinforcement of wing pattern (i.e., greater similarity) could occur when co‐mimics are in sympatry. We also examined variation in the red forewing band because this trait is critical for both mimicry and sexual communication. Morphometric results revealed significant but small differences among species, particularly in the shape of the forewing of co‐mimics. Although we did not observe greater similarity when co‐mimics were in sympatry, nearly identical patterns provided evidence of convergence for mimicry. In contrast, mimetic pairs could be distinguished based on the shape (but not the size) of the red band, suggesting an “advergence” process. In addition, sexual dimorphism in the red band shape (but not size) was found for all lineages. Thus, we infer that natural selection due to predation by birds might not be the only mechanism responsible for variation in color patterns, and sexual selection could be an important driver of wing phenotypic evolution in this mimicry ring.

Association between molecular markers and behavioral phenotypes in the immatures of a butterfly

Newly hatched caterpillars of the butterfly Heliconius erato phyllis routinely cannibalize eggs. In a manifestation of kin recognition they cannibalize sibling eggs less frequently than unrelated eggs. Previous work has estimated the heritability of kin recognition in H. erato phyllis to lie between 14 and 48%. It has furthermore been shown that the inheritance of kin recognition is compatible with a quantitative model with a threshold. Here we present the results of a preliminary study, in which we tested for associations between behavioral kin recognition phenotypes and AFLP and SSR markers. We implemented two experimental approaches: (1) a cannibalism test using sibling eggs only, which allowed for only two behavioral outcomes (cannibal and non-cannibal), and (2) a cannibalism test using two sibling eggs and one unrelated egg, which allowed four outcomes [cannibal who does not recognize siblings, cannibal who recognizes siblings, “super-cannibal” (cannibal of both eggs), and “super non-cannibal” (does not cannibalize eggs at all)]. Single-marker analyses were performed using χ2 tests and logistic regression with null markers as covariates. Results of the χ2 tests identified 72 associations for experimental design 1 and 73 associations for design 2. Logistic regression analysis of the markers found to be significant in the χ2 test resulted in 20 associations for design 1 and 11 associations for design 2. Experiment 2 identified markers that were more frequently present or absent in cannibals who recognize siblings and super non-cannibals; i.e. in both phenotypes capable of kin recognition.

Terpenoids dominate the bouquet of volatile organic compounds produced by Passiflora edulis in response to herbivory by Heliconius erato phyllis (Lepidoptera: Nymphalidae)

In response to injury, plants produce volatile organic compounds (VOCs) that usually differ depending on the type of damage they have suffered (e.g., mechanical damage, herbivory, and oviposition). The objectives of this study were to identify and compare the bouquet of volatiles emitted by passion vine plants (Passiflora edulis) after injury caused by mechanical damage (MD), herbivory (HB), and oviposition (OV) by the lepidopteran, Heliconius erato phyllis. Following injury, extracts of plant emissions were collected from each treatment every 24 h for three days and were analyzed by GC and GC/MS. Results show that plants emitted 12 volatiles before and after damage, namely terpenoids, ketones, and aldehydes. Although no significant differences were detected between the three treatments individually, if the entire bouquet of volatiles is analyzed, samples collected at 24 h were different from samples collected at 48 and 72 h. However, terpenoid emission increased significantly in HB plants after 24 h. HB plants emitted approximately 6300, 50, 46, 11, 6, and 3.6 times more (3E,7E)-4,8,12-trimethyl-1,3,7,11-tridecatetraene, (E)-4,8-dimethyl-1,3,7-nonatriene (DMNT), (E)-β-ocimene, (Z)-β-farnesene, (E)-β-caryophyllene, and farnesane, respectively, compared to control plants. OV plants displayed a peak of emission of (E)-β-ocimene after 72 h, which distinguished them from HB plants. MD plants showed a general increase of VOCs versus undamaged control plants. Furthermore, it has been suggested that (E)-β-ocimene may be sequestered by larvae of H. erato phyllis as a component of the odoriferous bouquet of the abdominal scent glands present in adult males, which play a role in sexual communication.

Dissecting the nature of subtle phenotypic variation in wing colour elements of Müllerian co-mimics

Abstract:Polymorphism is common in nature, but few Heliconius species are polymorphic for wing colour patterns. Eastern Brazil H. erato phyllis populations are polymorphic for hindwing elements (red raylets) and studies suggest that trait distribution varies seasonally. We carried a 3-y sampling to evaluate the hypothesis that season, wing length and pollen foraging were associated with morph diversity. Individual phenotypes were scored with regards to number of red elements in the dry and in the wet seasons. Co-mimic H. melpomene nanna was also analysed. We scored 432 H. erato and 513 H. melpomene. Our results confirm polymorphism in H. erato, with individuals showing from one to eight elements, with a mode between five and six. We found that H. melpomene is polymorphic for red dots, varying from two to five (mode = 2). Red basal dots were mostly invariant in H. erato. Even though we found a seasonal change in pollen loads, we found no association between individual phenotypes and season, pollen load scores, or wing length. We reject the hypothesis of ecological correlates of morph frequency, and suggest that trait colour variation in the two species is linked to and constrained by effects on mate recognition.

Comparative effects of inbreeding and outbreeding on the ontogeny ofHeliconius erato phyllis(Lepidoptera: Nymphalidae)

Inbreeding depression has been reported in various groups of organisms, including insects. Estimates of inbreeding consequences were obtained by comparing 12 life-history and morphological traits among nine inbred families (F = 0.25) and 16 outbred families (F = 0) of the Neotropical butterfly Heliconius erato phyllis. A Student's t-test showed statistically significant differences for pupal weight and right forewing area, both in males and in females, between inbred and outbred families. Survival during development, from egg hatching to adulthood, also differed significantly between inbred and outbred families. The average number of haploid lethal equivalents was 0.17 for pupal weight, 0.15 for forewing area and 0.71 for survival from hatching to adulthood. The results of this study confirm that the consequences of inbreeding are more deleterious to life history traits than to morphological ones.

Oviposition in Heliconius erato (Lepidoptera, Nymphalidae): how Essential Is Drumming Behavior for Host-Plant Selection?

Quantitative studies on the behavioral events involved in oviposition by phytophagous insects are scant. One of these events is drumming behavior, a rapid extension of forelegs after landing on a host, which remains largely under studied. In this study, quali- and quantitative analyses of Heliconius erato phyllis (Lepidoptera: Nymphalidae) oviposition events and drumming behavior in relation to both preferred (Passiflora misera and P. suberosa) and non-preferred (P. caerulea and P. alata) hosts were performed. For the first time, drumming behavior is described by images made during the butterfly oviposition process. The same set of females was assigned to both single- and multiple-choice tests under insectary conditions, and their behaviors toward each host were recorded on video. The resulting images were analyzed frame by frame. Both frequency and duration of behavioral events related to oviposition varied on preferred vs non-preferred hosts. On preferred hosts, behaviors that were associated with egg deposition (inspecting flight, drumming and abdomen touching) were pronounced; subsequently, oviposition occurred within a few seconds. On non-preferred host plants, behaviors that did not predict oviposition (flying and resting) predominated, and oviposition on these hosts was negligible. Drumming occurred on all plants and resulted in a faster decision-making process on high-quality hosts (prompt acceptance) as well as on lethal hosts (prompt rejection) compared to the host that confers poor larval performance (delayed rejection). Thus, drumming is crucial for decision-making related in host-plant selection by H. erato phyllis, and influences other behaviors involved in the oviposition process.

Kin recognition in a butterfly: inferences about its heritability

Some animal species are able to recognise their relatives, cannibalising mostly non-kin individuals, as happens with the caterpillars of the butterfly Heliconius erato phyllis. Here, we established a breeding programme where adults had different coefficients of relatedness (r = 0 and r = 0.5; offspring, respectively with F = 0 and F = 0.25). The caterpillars cannibalised or did not cannibalise sibling eggs, which was used to make inferences on the kin recognition inheritance of non-cannibals, and to estimate its heritability. The parents had both cannibal and non-cannibal phenotypes; crosses were done for all combinations of these phenotypes, which were divided into outbred and inbred. Caterpillars were grown in the laboratory until the adult stage, and were classified as cannibal or non-cannibal after a test with three sibling eggs. The results showed that the mean number of non-cannibal siblings, those that recognise their kin, is larger in inbreds, above 60%, regardless of their parents’ behaviour. Among non-inbred individuals the mean frequency of non-cannibal siblings was around 38%; these differences were statistically significant. The heritability was estimated by four methods. For F = 0 it was around 0.24 ± 0.11, and for F = 0.25 it was around 0.20 ± 0.17. These results, taken together, were discussed in terms of a threshold character: above this threshold the expression of the non-cannibal phenotype is allowed. As for the low values for heritabilities, they were discussed in terms of the theory of traits relevant to both fitness and the resource allocation priority hypothesis.

An experimental approach to test the source of information on kinship in butterfly immatures

This study focuses on a non-social species of butterfly of the Neotropical genus Heliconius, H. erato phyllis, where caterpillar-egg cannibalism may occur. In this species, newly hatched caterpillars are able to recognize sibling eggs, preferentially cannibalizing unrelated when there is a choice. The purpose of this study was to verify whether the information to recognize the egg as sibling or non-kin is in the chorion (maternal origin) and/or in the embryo/young caterpillar, by performing both caterpillar-egg and caterpillar-caterpillar cannibalism tests. The results of these tests suggest that the signal evaluated to recognize kinship is in the egg chorion, not in the caterpillar itself. We discuss these findings in the light of possible nutritional gain and the opportunity of encounters of immatures in a non-social butterfly.

Embryogenesis of Heliconius erato (Lepidoptera, Nymphalidae): a contribution to the anatomical development of an evo-devo model organism.

This study reports on the embryogenesis of Heliconius erato phyllis between blastoderm formation and the prehatching larval stage. Syncytial blastoderm formation occurred approximately 2 h after egg laying (AEL) and at about 4 h, the cellular blastoderm was formed. The germ band arose from the entire length of the blastoderm, and rapidly became compacted occupying approximately two-thirds of the egg length. At about 7 h AEL, protocephalon and protocorm differentiation occurred. Continued proliferation of the germ band was followed by penetration into the yolk mass, forming a C-shaped embryo at about 10 h. Approximately 12 h AEL, the gnathal, thoracic and abdominal segments became visible. The primordium of the mouthparts and thoracic legs formed as paired evaginations, while the prolegs formed as paired lobes. At about 30 h, the embryo reversed dorsoventrally. Approximately 32 h AEL, the protocephalon and gnathal segments fused, shifting the relative position of the rudimentary appendages in this region. At about 52 h, the embryo was U-shaped in lateral view and at approximately 56 h, the bristles began evagination from the larval cuticle. Larvae hatched at about 72 h. We found that H. erato phyllis followed an embryonic pattern consistent with long-germ embryogenesis. Thus, we believe that H. erato phyllis should be classified as a long-germ lepidopteran. The study of H. erato phyllis embryogenesis provided a structural glimpse into the morphogenetic events that occur in the Heliconius egg period. This study could help future molecular approaches to understanding the evolution of Heliconius development.

Defensive behavior associated with secretions from the prosternal paired glands of the larvae of Heliconius erato phyllis Fabricius (Lepidoptera, Nymphalidae)

Defensive behavior associated with secretions from the prosternal paired glands of the larvae of Heliconius erato phyllis Fabricius (Lepidoptera, Nymphalidae). Our work presents for the first time, the defensive behavior associated with the release of the product of the prosternal paired glands of the larva of Heliconius erato phyllis Fabricius, 1775 (Lepidoptera, Nymphalidae, Heliconiinae). The prosternal glands were first described for larvae of H. erato phyllis. They are formed by two types of glandular structures: the impair gland and the paired glands. The prosternal glands are located within the conical integumentary sac, which in turn is situated on the individual's prosternum. The main goal of this study is to analyze the existence of any secretion from the prosternal paired glands, and check the action mode of this secretion. The methodology used for chemical analysis of the glands included the aeration and, analysis in gas chromatography and gas chromatography-mass spectrometry. The results show that the prosternal glands do not produce volatiles. Bioassays were conducted with simulated and natural attacks and revealed that the prosternal paired glands produce secretions of defense together with silk produced by labials glands as a defense strategy, described for the first time, against ants. The strategy consists in wrapping the ant with silk threads, the entire wrapped object moved to the end of the body, with the aid of the legs and prolegs, and possibly fixed in a nearby place. Evidence for the existence of a conical integumentary sac in larvae of other species and families of Lepidoptera allows us to propose the possibility of occurrence of prosternal paired glands with defensive function in these other groups as well.

BUTTERFLIES OF CARIRI: PAPILIONOIDEA AND HESPERIOIDEA IN THE SLOPES OF THE CHAPADA DO ARARIPE IN CEARÁ, BRAZIL

This study aimed at conducting the first species inventory of butterflies in a location in the slopes of the Chapada do Araripe, in Ceara's Cariri region, in the Riacho do Meio Ecological Park. Specimens were collected from June 2009 to July 2010. We collected butterflies actively using insect nets, totaling 96 sampling hours. We collected a total of 236 individuals of 28 species and 5 families [Hesperiidae (N= 3), Lycaenidae (N= 1), Nymphalidae (N= 15), Pieridae (N= 7) and Riodinidae (N= 1)]. Nymphalidae was the most diverse family with 16 species and Heliconius erato phyllis (Fabricius) was the most abundant species.

Ultrastructure and morphogenesis of the wing scales in Heliconius erato phyllis (Lepidoptera: Nymphalidae): What silvery/brownish surfaces can tell us about the development of color patterning?

Usually the literature on Heliconius show three types of scales, classified based on the correlation between color and ultrastructure: type I – white and yellow, type II – black, and type III – orange and red. The ultrastructure of the scales located at the silvery/brownish surfaces of males/females is for the first time described in this paper. Besides, we describe the ontogeny of pigmentation, the scale morphogenesis and the maturation timing of scales fated to different colors in Heliconius erato phyllis. The silvery/brownish surfaces showed ultrastructurally similar scales to the type I, II and III. The ontogeny of pigmentation follows the sequence red, black, silvery/brownish and yellow. The maturation of yellow-fated scales, however, occurred simultaneously with the red-fated scales, before the pigmentation becomes visible. In spite of the scales at the silvery/brownish surfaces being ultrastructurally similar to the yellow, red and black scales, they mature after them; this suggests that the maturation timing does not show a relationship with the scale ultrastructure, with the deposition timing of the yellow pigment. The analysis of H. erato phyllis scale morphogenesis, as well as the scales ultrastructure and maturation timing, provided new findings into the developmental architecture of color pattern in Heliconius.

Oviposition preference and larval performance in a Heliconius erato phyllis (Lepidoptera: Nymphalidae) population from southeastern Brazil: is there a positive relationship?

Preference–performance hypothesis was tested in the butterfly Heliconius erato phyllis under field, insectary and laboratory conditions in a population from southeastern Brazil. Throughout a year, Passiflora capsularis was the host predominantly used for oviposition in the field. Similarly, when host plants were of high quality in the field, H. erato phyllis females that came from this environment laid more eggs on P. capsularis under insectary conditions. In contrast, when host plant quality declined in nature, females did not show preference for any host plant in particular in the insectary. Larvae had superior performances when reared on P. capsularis and Passiflora edulis than on Passiflora jilekii. Overall, our results show a trend towards a positive relationship between oviposition and performance in H. erato phyllis. The factors that might lead to an absence of selection under insectary conditions by females from a poor-quality environment are discussed.