Counterintuitive size patterns in bivoltine moths: late-season larvae grow larger despite lower food quality

Within a season, successive generations of short-lived organisms experience different combinations of environmental parameters, such as temperature, food quality and mortality risk. Adult body size of e.g. insects is therefore expected to vary both as a consequence of proximate environmental effects as well as adaptive responses to seasonal cues. In this study, we examined intraspecific differences in body size between successive generations in 12 temperate bivoltine moths (Lepidoptera), with the ultimate goal to critically compare the role of proximate and adaptive mechanisms in determining seasonal size differences. In nearly all species, individuals developing late in the season (diapausing generation) attained a larger adult size than their conspecifics with the larval period early in the season (directly developing generation) despite the typically lower food quality in late summer. Rearing experiments conducted on one of the studied species, Selenia tetralunaria also largely exclude the possibility that the proximate effects of food quality and temperature are decisive in determining size differences between successive generations. Adaptive explanations appear likely instead: the larger body size in the diapausing generation may be adaptively associated with the lower bird predation pressure late in the season, and/or the likely advantage of large pupal size during overwintering.     

Phenotypic plasticity in field populations of the tropical butterfly Hypolimnas bolina (L.) (Nymphalidae)

Phenotypic plasticity may enable organisms to maximize their fitness in seasonally variable environments. However, in butterflies, seasonal polyphenism is often striking but functionally obscure. This paper addresses the possible adaptive significance of phenotypic variation in the tropical butterfly Hypolimnas bolina (L.) (Nymphalidae). Plasticity in body size and wing coloration can be elicited in this species under laboratory conditions, however it is not known how this plasticity is expressed in the wild. Moreover, adult H. bolina spend the winter dry season in a reproductive diapause, which allows certain predictions regarding the occurrence of seasonal plasticity. Based on consideration of the requirements of diapausing and directly developing individuals, we predicted that if seasonal plasticity in phenotype were adaptive, then overwintering individuals should be larger and darker than their directly developing counterparts. This prediction was largely - although not entirely - fulfilled. Dry season butterflies were duller and darker than their wet season counterparts (this plasticity was superimposed on a genetic colour polymorphism), however size plasticity varied geographically. Dry season adults were consistently larger than wet season adults in the tropical north, but not in the south. We use these findings to discuss the possible adaptive significance of seasonal variation in the colour and size of this tropical butterfly.     

Northward range expansion requires synchronization of both overwintering behaviour and physiology with photoperiod in the invasive Colorado potato beetle (Leptinotarsa decemlineata)

Photoperiodic phenological adaptations are prevalent in many organisms living in seasonal environments. As both photoperiod and growth season length change with latitude, species undergoing latitudinal range expansion often need to synchronize their life cycle with a changing photoperiod and growth season length. Since adaptive synchronization often involves a large number of time-consuming genetic changes, behavioural plasticity might be a faster way to adjust to novel conditions. We compared behavioural and physiological traits in overwintering (diapause) preparation in three latitudinally different European Colorado potato beetle (Leptinotarsa decemlineata) populations reared under two photoperiods. Our aim was to study whether behavioural plasticity could play a role in rapid range expansion into seasonal environments. Our results show that while burrowing into the soil occurred in the southernmost studied population also under a non-diapause-inducing long photoperiod, the storage lipid content of these beetles was very low compared to the northern populations. However, similar behavioural plasticity was not found in the northern populations. Furthermore, the strongest suppression of energy metabolism was seen in pre-diapause beetles from the northernmost population. These results could indicate accelerated diapause preparation and possibly energetic adjustments due to temporal constraints imposed by a shorter, northern, growth season. Our results indicate that behavioural plasticity in burrowing may have facilitated initial range expansion of L. decemlineata in Europe. However, long-term persistence at high latitudes has required synchronization of burrowing behaviour with physiological traits. The results underline that eco-physiological life-history traits of insects, such as diapause, should be included in studies on range expansion.     

Adaptive latitudinal cline of photoperiodic diapause induction in the parasitoid Nasonia vitripennis in Europe

Living in seasonally changing environments requires adaptation to seasonal cycles. Many insects use the change in day length as a reliable cue for upcoming winter and respond to shortened photoperiod through diapause. In this study, we report the clinal variation in photoperiodic diapause induction in populations of the parasitoid wasp Nasonia vitripennis collected along a latitudinal gradient in Europe. In this species, diapause occurs in the larval stage and is maternally induced. Adult Nasonia females were exposed to different photoperiodic cycles and lifetime production of diapausing offspring was scored. Females switched to the production of diapausing offspring after exposure to a threshold number of photoperiodic cycles. A latitudinal cline was found in the proportion of diapausing offspring, the switch point for diapause induction measured as the maternal age at which the female starts to produce diapausing larvae, and the critical photoperiod for diapause induction. Populations at northern latitudes show an earlier switch point, higher proportions of diapausing individuals and longer critical photoperiods. Since the photoperiodic response was measured under the same laboratory conditions, the observed differences between populations most likely reflect genetic differences in sensitivity to photoperiodic cues, resulting from local adaptation to environmental cycles. The observed variability in diapause response combined with the availability of genomic tools for N. vitripennis represent a good opportunity to further investigate the genetic basis of this adaptive trait.     

Expression of alternative developmental pathways in the cabbage butterfly,Pieris melete and their differences in life history traits

The seasonal life cycle of the cabbage butterfly, Pieris melete is complicated because there are three options for pupal development: summer diapause, winter diapause, and nondiapause. In the present study, we tested the influence of temperature, day length, and seasonality on the expression of alternative developmental pathways and compared the differences in life history traits between diapausing and directly developing individuals under laboratory and field conditions. The expression of developmental pathway strongly depended on temperature, day length, and seasonality. Low temperatures induced almost all individuals to enter diapause regardless of day length; relatively high temperatures combined with intermediate and longer day lengths resulted in most individuals developing without diapause in the laboratory. The field data revealed that the degree of phenotypic plasticity in relation to developmental pathway was much higher in autumn than in spring. Directly developing individuals showed shorter development times and higher growth rates than did diapausing individuals. The pupal and adult weights for both diapausing and directly developing individuals gradually decreased as rearing temperature increased, with the diapausing individuals being slightly heavier than the directly developing individuals at each temperature. Female body weight was slightly lower than male body weight. The proportional weight losses from pupa to adult were almost the same in diapausing individuals and in directly developing individuals, suggesting that diapause did not affect weight loss at metamorphosis. Our results highlight the importance of the expression of alternative developmental pathways, which not only synchronizes this butterfly's development and reproduction with the growth seasons of the host plants but also exhibits the bet‐hedging tactic against unpredictable risks due to a dynamic environment.     

Life history plasticity in the satyrine butterfly Lasiommata petropolitana: investigating an adaptive reaction norm

This study addresses the general hypothesis that insects living in seasonal environments should shorten development times at progressively later dates in the growth season, and that insects living outside equatorial areas should use daylength as a cue to determine the date. Diapause strategies and reaction norms relating the duration of larval development to daylength was investigated in a French population of the butterfly, Lasiommata petropolitana. The results are compared with those of an earlier study of the species in Sweden. Because of the diapausing strategy and phenology of the population, it was expected that an adaptive reaction norm relating larval time to daylength should have a positive slope, i.e. relatively shorter daylengths induce faster growth and development. This prediction was supported, and the reaction norm was qualitatively similar to the one found in Swedish populations. In the French population it was, however, shifted to a range of shorter photoperiods which corresponds to the regime of shorter daylengths in southern Europe. Shorter larval development times and high growth rates were associated with a reduction in pupal size, suggesting a trade off between time and size at pupation. There was no evidence of a trade off between growth rate and starvation endurance. The results suggests that the daylength-dependent decision of what growth trajectory an individual larva will follow, is not made continuously but rather at one or a few occasions during larval development. It is clear that larvae of L. petropolitana make developmental decisions in relation to the daylength they experience during larval growth. The result is a reaction norm that agrees closely to what is predicted by some life history models, suggesting that it is an adaptation for optimising life history traits in a seasonal environment.     

The environmental and genetic control of seasonal polyphenism in larval color and its adaptive significance in a swallowtail butterfly

Seasonal polyphenism, in which different forms of a species are produced at different times of the year, is a common form of phenotypic plasticity among insects. Here I show that the production of dark fifth-instar caterpillars of the eastern black swallowtail butterfly, Papilio polyxenes, is a seasonal polyphenism, with larvae reared on autumnal conditions being significantly darker than larvae reared on midsummer conditions. Both rearing photoperiod and temperature were found to have individual and synergistic effects on larval darkness. Genetic analysis of variation among full-sibling families reared on combinations of two different temperatures and photoperiods is consistent with the hypothesis that variation in darkness is heritable. In addition, the genetic correlation in larval darkness across midsummer and autumnal environments is not different from zero, suggesting that differential gene expression is responsible for the increase in larval darkness in the autumn. The relatively dark autumnal form was found to have a higher body temperature in sunlight than did the lighter midsummer form, and small differences in temperature were found to increase larval growth rate. These results suggest that this genetically based seasonal polyphenism in larval color has evolved in part to increase larval growth rates in the autumn.     

Seasonal plasticity in life history traits: growth and development in Polygonia c-album (Lepidoptera: Nymphalidae)

The potentially multivoltine comma butterfly, Polygonia c-album L., hibernates in the adult stage. The adult seasonal morph is demonstrated to be a good indicator of whether an individual has entered reproductive diapause or is developing directly to sexual maturation. This fact, and the assumption that a short development time is not equally important to all categories of individuals, was used to test predictions on variation in life-history traits among categories (morphs and sexes) and environments (temperature and photoperiod) at the level of individuals and to some extent families and populations (the univoltine Stockholm population and the partially bivoltine Oxford population). Individuals developing to adults in a short time were expected to be smaller and lighter as a result of a basic trade-off between the two traits. Development times varied in accordance with predictions, but in most cases this was due to plastic growth and development in both the larval and pupal stages rather than through variation in size or weight, i.e. size was a highly canalized trait. This suggests a relationship between plasticity and canalization and a strong potential for plasticity to shield life-history traits from selection. Individuals regulated development times also within developmental pathways, in response to photoperiods indicating the progression of the season. These and other results suggest that development times are not normally minimized in temperate butterflies unless this is enforced by direct development and protandry. There is thus scope for a high degree of adaptive plasticity in growth- and developmental rates which may devalue the basic trade-offs assumed by life-history theory and account for inconsistencies with its predictions.     

Seasonal plasticity in growth and development of the speckled wood butterfly, Pararge aegeria (Satyrinae)

Regulation of growth and development by photoperiod was studied in a population of the speckled wood butterfly, Purarge aegeria L. (Lepidoptera: Satyrinae), from southern Sweden. Individuals were reared in a range of photoperiodic regimes (9L. to 22L) and temperatures (13°C to 21° C). Plasticity was found for important life-history traits- generation time, growth rate and final weight and seasonal regulation of development in response to photoperiod was found to occur at two levels. Purarge aegeria hibernates as a third instar larva or in the pupal stage, cantering one of four major developmental pathways in response to photoperiod: (1) direct development in both the larval and pupal stages, (2) pupal winter diapause with or (3) without a preceding larval summer diapause, or (4) larval winter diapause. In addition to this high-level regulation of individual development, larval growth rate and pupal development rate also appear to be finally regulated by photoperiod within each major pathway. As photoperiods decreased from 22 h to 17 h at 17° C, growth rate among directly developing larvae increased progressively, as was the case for larva? developing according to a univoltine life cycle from 17 h to 14 h. At two photoperiods, 13 h and 16 h (corresponding to shifts between major pathways), both larval and pupal development were extremely variable with the fastest individuals developing directly and the slowest developing with a diapause. This indicates a gradual nature of diapause itself, suggesting that the two level may not he fundamentally different.     

Changes of biogenic amine levels in haemolymph during diapausing and non-diapausing status in Pieris brassicae L

During the larval development of Pieris brassicae, photoperiod status induces either direct (non-diapausing) development or diapausing development. Biogenic amines such as dopamine and serotonin and their derivatives may be implicated in the physiological control and adaptation of these insects to different photoperiods. High performance liquid chromatography coupled with electrochemical detection was used to measure biogenic amine concentrations in insect haemolymph in different photoperiod conditions. At the end of the higher photosensitivity phase, dopamine levels were two-fold higher in diapausing (0.914 pmol/microl+/-0.025) as compared to non-diapausing insects (0.415 pmol/microl+/-0.02; P<0.001). Globally, dopamine catabolites are found in higher levels in non-diapausing insects (in prepupa, near seven times more for NADA, and 25 times more for DOPAc), thus indicating a higher dopamine turn-over. Serotonin levels were detected only at the beginning of this instar in diapausing insects but not in direct development insects. During pupal life, dopamine levels were significantly higher in diapausing than in non-diapausing insects (in 3-day pupae, dopamine was 28 times higher), and remained at high levels throughout diapausing life. At the beginning of diapause, serotonin levels were higher in diapausing insects than in non-diapausing insects (in 3-day pupae, serotonin was 13 times higher). These levels decreased in the middle of diapause period, and then increased just before its breaking. Dopamine and serotonin levels presented the typical kinetics of diapause development and may be used as an indicator of the evolution of diapause status.     

Influence of seasonal time constraints on growth and development of common frog tadpoles: a photoperiod experiment

Organisms living in seasonal environments are often limited by the time available to complete their development. Especially individuals in northern populations may face severe time constraints in their need of completing development before the end of the growth season. Larval amphibians have been widely used in studies of phenotypic plasticity. However, their responses to changes in photoperiod, the main seasonal cue in many organisms, are unknown. In a laboratory experiment, we studied whether common frog (Rana temporaria) tadpoles originating from two populations (separated latitudinally by 1600 km) adjust their growth and development according to the progress of the season by using photoperiodic cues, and whether these responses are temperature dependent. We hypothesised that if frogs use photoperiod as a cue, they should increase growth and development rates as a response to photoperiodic treatments mimicking progressing season. Although our predictions were not verified in either of the populations, photoperiod manipulations had effects on larval life history in both populations. When exposed to progressing season treatments and high temperature, tadpoles from the southern population ceased feeding, which led to delayed metamorphosis and increased mortality. In the northern population, age at metamorphosis was unaffected by the photoperiod treatments, but growth rate until metamorphosis and metamorphic size were slightly larger in the treatments with shorter (increasing or decreasing) day length. Irrespective of photoperiod treatments, growth and development rates, size at metamorphosis and food consumption were higher in the northern as compared to the southern population. These results indicate that in contrast to several insect species, the critical life history decisions in amphibian larvae may not be strongly influenced by photoperiodic cues, but different populations seem to differ in this respect. However, the strong temperature×photoperiod interactions in several traits in the southern population suggest that the role of photoperiodic cues may be affected by other environmental factors, although the ecological significance of these differences remains unclear.     

Effects of thermophotoperiod on growth parameters of Helicoverpa armigera

The effects of photoperiod and temperature on growth parameters of Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae) were investigated under laboratory conditions by exposing its larvae to fluctuating and alternating temperature regimes. Our data made evident that the interplay between photoperiod and temperature has a direct effect on growth parameter responses of H. armigera. However, the type of temperature regime (fluctuating or alternating) may enhance or diminish the effects of photoperiod. With fluctuating temperature, larval developmental time was significantly reduced with increasing photophase, irrespective of the diapause status later in the pupal stage; the lowest value was observed under continuous light conditions. With alternating temperature, larval developmental time was significantly decreased with the reduction in temperature amplitude, and with the coincidence of the thermophase with the scotophase. At both temperature regimes, no significant differences in pupal weight were observed between diapausing and non‐diapausing individuals, although, in most treatments, the diapausing pupae tended to be heavier than the non‐diapausing ones. The increased pupal weight of individuals destined for diapause is probably related to their longer larval developmental time. Knowing the effects of these factors on the variation of H. armigera growth is necessary in explaining phenomena associated with immature stages, and can lead to a more profound understanding of the potential for this insect to evolve in response to environmental changes.     

Architectural plasticity in a Mediterranean winter annual

Size variability in plants may be underlain by overlooked components of architectural plasticity. In annual plants, organ sizes are expected to depend on the availability and reliability of resources and developmental time. Given sufficient resources and developmental time, plants are expected to develop a greater number of large branches, which would maximize fitness in the long run. However, under restrictive growth conditions and environmental reliability, developing large branches might be risky and smaller branches are expected to foster higher final fitness. Growth and architecture of Trifolium purpureum (Papilionaceae) plants from both Mediterranean (MED) and semi-arid (SAR) origins were studied, when plants were subjected to variable water availability, photoperiod cues and germination timing. Although no clear architectural plasticity could be found in response to water availability, plants subjected to photoperiod cuing typical to late spring developed fewer basal branches. Furthermore, plants that germinated late were significantly smaller, with fewer basal branches, compared with plants which grew for the same time, starting at the beginning of the growing season. The results demonstrate an intricate interplay between size and architectural plasticities, whereby size modifications are readily induced by environmental factors related to prevalent resource availability but architectural plasticity is only elicited following the perception of reliable anticipatory cues.     

Critical photoperiod and daylength threshold differences between northern and southern populations of the butterfly Limenitis archippus

Laboratory studies show that different photoperiods induce diapause in northern (Vermont) and southern (Maryland) larval strains of the butterfly Limenitis archippus. The northern strain responds to $\text{1}\text{2}\text{hr}$ longer photoperiod thresholds and critical ranges than does the southern one. These responses are correlated with geographic differences in the ambient photoperiod of the two localities. In this facultative diapausing species, third instar larvae construct hibernacula within the basal portions of tubular leaves spun with silk, when daylength approaches either 13·5 hr (Vermont strain) or 13·0 hr (Maryland strain). When reared in total darkness some larvae develop directly to fourth inszar without diapause, although mortality is high. Among both strains different broods exhibit different incidences of diapause. Reciprocal inter-strain hybrids show intermediate diapause responses, suggesting that larval diapause is under the control of multiple genes.     

Temperature- and density-dependence of diapause induction and its life history correlates in the geometrid moth Chiasmia clathrata (Lepidoptera: Geometridae)

The relative roles of genetics and developmental plasticity in creating phenotypes adapted to prevailing conditions are insufficiently understood. In potentially multivoltine temperate insects, individuals that do not enter diapause but develop directly into reproductive adults within the same season are severely time-constrained. Direct development is, however, under selection only if expressed in the wild. Thus, adaptive correlates of the direct development are expected to evolve and persist only in multivoltine populations. We studied the genetic and phenotypic components of variation in juvenile development in the geometrid moth Chiasmia clathrata from univoltine and bivoltine regions. Larvae were reared at two temperatures (14/20 °C) and densities (low/high) in a factorial split-brood experiment. High temperature and low density promoted direct development, the former condition being associated with a short development time, high growth rate and large body size. Genotypes of bivoltine origin had a higher propensity for direct development and seemingly expressed an exaggerated plastic response to increasing temperature compared to the ones from univoltine populations. Alternative life history phenotypes associated with the induced developmental pathway emerged only in the bivoltine region, direct development resulting in a short larval period, high growth rate and small size at 20 °C there. The degree of differentiation between the developmental pathways was insensitive to larval density; high density only decreased both development time and body size to a certain degree. We conclude that the differences between the pathways are not due to the induction of a particular pathway itself, but geographically varying selection pressures shape the correlation structure among life history traits and their pathway-specific expression.     

Developmental response to a seasonal time constraint: the effects of photoperiod on reproduction in the grasshopper Romalea microptera

Abstract.  1. Some organisms respond adaptively to seasonal time constraints by altering development time to life-history transitions (e.g. metamorphosis, oviposition). Such life-history changes may have costs (e.g. reduced fecundity, mass, offspring quality).
2. The hypothesis that a northern population of the grasshopper Romalea microptera (Beauvois) would show adaptive plasticity in oviposition timing in response to seasonal time constraints was tested by manipulating photoperiod to simulate the middle of the active season (Long photoperiod), the end of the active season (Short photoperiod), and seasonal change (photoperiod Declining from long to short). Females received either high or low food rations. Short or Declining photoperiod were predicted to induce early oviposition with costs of reduced egg number, post-oviposition mass, or egg size, particularly in low-food females.
3. Effects of food ration and photoperiod, but not interaction, were significant in failure time analysis of age at oviposition. mancova on age at oviposition, egg number, and post-oviposition mass yielded similar effects. The multivariate effect of photoperiod resulted primarily from reduced time to oviposition in Short or Declining photoperiod. No costs in egg number or post-oviposition mass were associated with this photoperiod-induced reduction in time to oviposition. The multivariate effect of food ration resulted mainly from lower egg number with low food. Food ration affected egg size, but photoperiod and interaction did not. In all cases, Short and Declining photoperiod produced similar effects.
4. In its northern range, R. microptera accelerates reproduction in response to seasonal constraints, a response that may be adaptive. How R. microptera avoids costs associated with this reduced pre-oviposition period remains unknown.     

Effect of photoperiod and temperature on diapause in a Florida strain of the tropical warehouse moth Ephestia cautella

A photoperiodically-controlled diapause of the long-day, short-day type was identified in a brown-winged, yellow-eyed strain of Ephestia cautella (Walker). The proportion of larvae diapausing in very long photoperiods was less than in short photoperiods. The mean critical photoperiod, here defined as that photoperiod giving half the maximum percentage of insects that diapause in response to photoperiod at a given temperature, was between 12 and 13 hr for the long-day reaction at both 20 and 25°C. The principal sensitive phase occurred near the time of the last larval moult. The mean duration of diapause was 2–3 months at 20°C and slightly longer at 25°C. The optimum temperature for diapause development was near 15°C, all larvae pupating within 24 days after a 45-day exposure at this temperature. Diapause could be terminated whenever larvae diapausing at 20°C were exposed to as few as five long (15 hr) photoperiods at 25°C. Long photoperiods at 20°C, or short photoperiods (9 hr) at 25°C were less effective in terminating diapause.     

Induction of diapause and seasonal morphs in butterflies and other insects: knowns,unknowns and the challenge of integration

The ‘choice’ of whether to enter diapause or to develop directly has profound effects on the life histories of insects, and may thus have cascading consequences such as seasonal morphs and other less obvious forms of seasonal plasticity. Present knowledge of the control of diapause and seasonal morphs at the physiological and molecular levels is briefly reviewed. Examples, mainly derived from personal research (primarily on butterflies), are given as a starting point with the aim of outlining areas of research that are still poorly understood. These include: the role of the direction of change in photoperiod; the role of factors such as temperature and diet in modifying the photoperiodic responses; and the role of sex, parental effects and sex linkage on photoperiodic control. More generally, there is still a limited understanding of how external cues and physiological pathways regulating various traits are interconnected via gene action to form a co‐adapted complete phenotype that is adaptive in the wild despite environmental fluctuation and change.     

Growth strategies and optimal body size in temperate pararginii butterflies

In temperate insects the evolution of growth strategies andthe optimal age and size at maturity will depend strongly onseasonal variation in temperature and other resources. However,compared to photoperiod, temperature itself is a relativelypoor predictor of seasonal change and timing decisions in insectsare often most strongly influenced by the photoperiod. HereI review the evolution of seasonal growth strategies in thebutterfly tribe Pararginii (Satyrinae: Nymphalidae) and relateit to life history theory. The results indicate that individuallarvae may adjust their growth trajectories in relation to informationon time horizons obtained from the photoperiod. The growth strategiescan be characterized by a set of state-dependent decision rulesthat specify how an individual should respond to its internalstate and external circumstances. These decision rules may alsoinfluence how individual growth change with a rise in temperature,showing that the standard expectation of increased growth rateswith increasing temperatures may not always be true. With lesstime available individual larvae increase growth rates and therebyachieve shorter development times, most often without any effectson final sizes. One reason for the apparent optimization ofgrowth rate seems to be that growing fast may incur costs thatlarvae developing under lower time limitations chose to avoid.The patterns of growth found in these and many other studiesare difficult to reconcile with common assumptions of what typicallydetermines optimal body size in insects. In particular it seemsas if there should be some costs of a large body size that,so far, have been poorly documented.     

Pupal diapause of Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae) mediated by larval host plants: pupal weight is important

Facultative diapause, a strategy that allows insects to initiate additional generations when conditions are favorable or to enter diapause when they are not, has a profound effect on the ecology and evolution of species. Most previous studies have concentrated on the role of photoperiod and temperature in inducing facultative diapause in insects. In contrast, here we studied pupal diapause mediated by larval host plants in the cotton bollworm Helicoverpa armigera, and confirmed that pupal weight is a critical factor. Two groups of third instar H. armigera larvae, kept at 25 °C with L:D = 8:16 and 20 °C with photoperiod of L:D = 8:16, respectively, were fed on six host plants and on artificial diet (as a control) to determine how larval host plants affect diapause incidence and related traits (such as pupal weight and developmental duration). The data showed larval host plants affected diapause incidence significantly and the effects could be masked by low temperature. Further analysis showed that pupal size, not the length of the sensitive stage, affected the decision to enter diapause. In a further experiment, third-instar to final-stage larvae deprived of artificial diet for 2 days demonstrated a direct relationship between pupal weight and diapause incidence. These results suggest that larval host plants, by affecting pupal size, may influence diapause occurrence in H. armigera. This has important adaptive significance for both over-wintering survival and the possibility for completing an additional generation.