Variability of photosensitive period and voltinism among populations of a butterfly,Ypthima multistriata,inhabiting similar latitudes and altitudes

In Ypthima multistriata Butler (Lepidoptera: Nymphalidae), there are univoltine and bivoltine populations in adjacent areas with similar climatic conditions. A previous study revealed that larvae of both univoltine and bivoltine populations diapause under a constant short day (i.e., a constant short light period; L13:D11), but not under a constant long‐day condition (L16:D8). However, in both types of populations, adults of an overwintering generation appear and oviposit in June and soon thereafter larvae hatch. Therefore, the younger larvae (at least the first instars) of both types of populations experience a long day; nevertheless, the larvae of univoltine populations diapause in nature. To resolve this inconsistency, we set up two hypotheses: (1) the photosensitive stage of larvae is the second instar or later, and (2) the photosensitive stage of univoltine populations is later than that of bivoltine populations. To test these hypotheses, we performed rearing experiments with two univoltine populations and two bivoltine ones. The results indicated that the photosensitive stage was the second or third instar and that the photosensitive stage was later in one univoltine population than in the two bivoltine populations. Larvae of the other univoltine population diapaused under all conditions. The former result supports our hypothesis, and the latter result indicates that the response to photoperiod is different among univoltine populations. In addition, larval development was slower in one univoltine population than in the bivoltine populations, which also delays the timing of the diapause decision in this univoltine population. Larvae that experienced a long day during the first and middle instars but experienced a short day at the end of their larval stage developed faster than larvae that experienced a constant long day. This may be an adaptation to enable emergence before the start of a cold season that is unsuitable for reproduction.     

A population model for diapausing multivoltine insects under asymmetric cannibalism

Pupa-eating cannibalism occurs naturally in several insect species. Byasa alcinous is a multivoltine species of Red-bodied Swallowtail butterfly found in East Asia, which diapauses as pupa over the winter and whose larvae cannibalize eggs and pupae. We investigate the effects on population dynamics of increasing the asymmetric cannibalistic attack rate of a general insect species in different environmental conditions. We do this by theoretically formulating a generalized system of univoltine and bivoltine larvae over two generations in the spring and summer months. We predict that a lack of resources over the summer can force the population to become entirely univoltine, unless the second-generation bivoltine larvae increase their cannibalistic attack rate, and consume the diapausing pupae from the first generation. The model shows that under extreme environmental conditions, the persistence of univoltine larvae is favoured when faced with the threat of extinction. The model also predicts the conditions for the coexistence of both univoltine and bivoltine larvae, and the degree to which they can both coexist, which decreases as the resource in the second generation increases. This work provides the grounding for future theoretical and experimental consideration of the role of cannibalism in determining insect voltinism.     

Optimal seasonal timing of univoltine and bivoltine insects

This study examines the optimal seasonal timing of the life cycle for univoltine and bivoltine insects, assuming that resource availability has a peak in the middle of a year and is symmetric around it. Results show that if the growth rate increases in proporrion to the bodyweight, bivoltine life cannot be optimal. If the growth rate is a power function of the bodyweight with a power smaller than unity, a symmetric bivoltine solution can be the optimal provided that the resource availability has a plateau in the middle of the season. If the resource availability has a sharp peak, the optimal pattern is an asymmetric bivoltine solution in which the larval periods of two generations differ in length. The bivoltine life cycle is more likely to be superior to the univoltine one if: growth is fast, suitable growing season is long, biomass loss during nonlarval stages is small, and egg size is small.     

亚洲玉米螟化性与繁殖力的关系

通过对发生在吉林省的一、二化性玉米螟的繁殖能力研究得出,不同化性越冬代玉米螟的产卵期、成虫寿命、单雌产卵量和最大产卵量差异显著,一化性越冬代玉米螟的产卵量比二化性玉米螟高４５％以上。经虫体大小和重量比较得出,一化性越冬代玉米螟的蛹重比二化性玉米螟的蛹重高３５％以上,人工饲养的一化性玉米螟Ｆ_1代的蛹重也比二化性的蛹重高１０％左右。     

Diapause and diapause dynamics of Colias alexandra (Lepidoptera: Pieridae)

Summary The mechanisms and adaptations involved in the obligate third instar larval diapause of Colias alexandra, a native Rocky Mountain and intermountain region butterfly, are examined. Generally univoltine throughout its distribution, scattered, isolated bivoltine populations occur. Factors influencing continuous development were investigated under laboratory conditions on a single generation of offspring from a univoltine population each year from 1976–1979. If exposed to mean temperatures >24°C during the second instar, a significant number of larvae fail to diapause. These results are interpreted in the context of the actual environmental conditions experienced by the univoltine source population and a bivoltine population. A high degree of individual variability in response to continuous development conditions is shown. Variation in certainty of diapause, within an originally univoltine population, could be the evolutionary starting point for evolving a genuine multivoltine cueing system and provide the mechanism for ecological range expansion.     

Effect of variation in photoperiodic response on diapause induction and developmental time in the willow leaf beetle, Plagiodera versicolora

The willow leaf beetle, Plagiodera versicolora (Coleoptera: Chrysomelidae) overwinters in adult diapause. In this study, the photoperiodic responses for diapause induction and developmental time were examined in the Ishikari (Hokkaido, Japan) population of P. versicolora. All females entered reproductive diapause under short daylength (L10:D14), but 31.7% of females did not enter diapause under long daylength (L16:D8). The developmental time from oviposition to adult emergence was significantly longer at L10:D14 than that at L16:D8. Norm of reaction curves illustrated variation among families in the photoperiodic responses for diapause induction and for developmental time. ANOVA indicated significant family × photoperiod interactions in the developmental time. At L16:D8, developmental time was positively correlated with the incidence of diapause in females. This means that a female having a longer developmental time tends to have a longer critical photoperiod. Such variation may be maintained by differences in selection pressures on the growth rate and the critical photoperiod for diapause induction between univoltine and bivoltine genotypes because Ishikari is located in a transitional area between populations with univoltine and bivoltine life cycles.     

Geographical variation in insect developmental period: effect of host plant phenology on the life cycle of the bruchid seed feeder shape Kytorhinus sharpianus

The voltinism of the bruchid Kytorhinus sharpianus Bridwell (Coleoptera: Bruchidae) and the phenology of its host plant Sophora flavescens Aiton (Leguminosae) were observed at four latitudes: Aomori (40°46' N), Obanazawa (38°37' N), Kujiranami (37°21' N) and Mitsuma (36°05' N) in northeastern Honshu (Japan). Kytorhinus sharpianus life cycle ranged from bivoltine and partially trivoltine in the south to univoltine and partially bivoltine in the north. Sophora flavescens started growing later in spring at higher latitudes. However, the relative growth rate was higher in the north (Aomori) than in the south (Mitsuma). In parallel with this, the first-generation of adult K. sharpianus appeared later at higher latitudes. When the four local populations were reared at 24 °C, L16:D8 and 65% r.h., males developed faster than females. The mean developmental time showed a saw-toothed latitudinal cline. The reversion in the latitudinal trend of variation corresponded to the change in the major type of life cycles from univoltine to bivoltine. Two heat units throughout the year and post-fruiting period were calculated as the sums of degree-days above the developmental threshold (12 °C) of K. sharpianus. Both heat units decreased in parallel with each other with increasing latitude. The greater growth rate of hosts in the northern population compensated for the smaller heat units. In addition, when the heat units were divided by the degree-days needed to complete development, the numerical value was the approximate number of generations observed in each locality.     

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.     

Latitudinal clines in alternative life histories in a geometrid moth

The relative roles of genetic differentiation and developmental plasticity in generating latitudinal gradients in life histories remain insufficiently understood. In particular, this applies to determination of voltinism (annual number of generations) in short‐lived ectotherms, and the associated trait values. We studied different components of variation in development of Chiasmia clathrata (Lepidoptera: Geometridae) larvae that originated from populations expressing univoltine, partially bivoltine or bivoltine phenology along a latitudinal gradient of season length. Indicative of population‐level genetic differentiation, larval period became longer while growth rate decreased with increasing season length within a particular phenology, but saw‐tooth clines emerged across the phenologies. Indicative of phenotypic plasticity, individuals that developed directly into reproductive adults had shorter development times and higher growth rates than those entering diapause. The most marked differences between the alternative developmental pathways were found in the bivoltine region suggesting that the adaptive correlates of the direct development evolve if exposed to selection. Pupal mass followed a complex cline without clear reference to the shift in voltinism or developmental pathway probably due to varying interplay between the responses in development time and growth rate. The results highlight the multidimensionality of evolutionary trajectories of life‐history traits, which either facilitate or constrain the evolution of integrated traits in alternative phenotypes.     

Changes in voltinism in a pine moth <Emphasis Type="Italic">Dendrolimus spectabilis</Emphasis> (Lepidoptera: Lasiocampidae) population: implications of climate change

Climate change induces an alteration in the life cycle of many poikilothermic organisms, resulting in changes in the structure and function of communities. Changes in voltinism in the pine moth Dendrolimus spectabilis (Butler), which is known to be univoltine in South Korea, were studied to elucidate the effects of climate change on their voltinism. The developmental stages of the pine moth were evaluated through field surveys, and the developmental rate was estimated at five different temperatures: 17, 20, 25, 28, and 32°C. Field surveys showed that the moths completed two generations per year, indicating that the phenology of the pine moth in this area had changed from univoltinism to bivoltinism. Laboratory experiments showed that increasing the temperature could induce a change in voltinism in the pine moth population. Generations of the bivoltine population displayed phenotypic plasticity: the fitness of the first generation was greater than that of the second generation with regard to size and fecundity. The difference in fitness between the first and second generations could be due to the influence of factors such as low food quality and heat stress on the second generation. Therefore, changes in thermal conditions due to climate change have offered this species the chance to develop a bivoltine population, but they have also exerted ecological costs, especially for the second generation of the pine moth.     

Life-history strategies and production of Tinodes waeneri (L.) (Trichoptera) in Lake Esrom,Denmark

The life cycle and production of Tinodes waeneri (L.) was examined at five shallow littoral localities in Lake Esrom, Denmark, during 1979/80. Five larval instars were demonstrated. The differences in temporal composition of instars indicated three different life cycles. A bivoltine strategy was shown for the most exposed locality which also had the highest average abundance equal to 11 500 ind. m^–2. A univoltine population was found in the more sheltered NW part of the lake with an average density of 1 500 ind. m^–2. At the three remaining stations the life cycles were partly bivoltine with wintering populations of mixed cohorts and densities between 10 500 and 3 000 ind. m^–3. Two distinct flight periods with maxima in June and August were demonstrated for bi- and partly bivoltine populations. Differences in rates of growth and elimination added to the variability in life cycle. Growth rates peaked in August (5.0–7.4% d^–1) at an average temperature of 20°C. Estimates of production ranged from 1.9 to 17.5 g AFDW m^–2 a^–1 with an average of 8.0 g AFDW m^–2. The P/B ratios, which were within the expected range for uni- and bivoltine populations of trichopterans, were 5.73 for the bivoltine population and 3.31 for the univoltine population.Estimates of consumption revealed that the populations could be sustained by autochthonous epilithic production in the habitats. Variability in the quality and seasonal availability of food were judged to influence the type of life cycle.Lake Esrom Littoral Research Publ. No. 8. Publication No. 385 from the Freshwater Biological Laboratory, University of Copenhagen, 51, Helsingørsgade, DK-3400 Hillerød, Denmark.Lake Esrom Littoral Research Publ. No. 8. Publication No. 385 from the Freshwater Biological Laboratory, University of Copenhagen, 51, Helsingørsgade, DK-3400 Hillerød, Denmark.     

Ecological and genetic correlates of range expansion in Coenonympha tullia

Range expansion by the North American butterfly species Coenonympha tullia is associated with dramatic changes in life history and in genetic and morphological variation. Eight of ten independent, variable loci exhibit step-clines in allele frequency; step-clines also occur in four wing pattern characters. Populations from the old part of the range are univoltine, and have significantly less genetic and morphological variation than populations from the recently colonized ranger, which arc bivoltine. The concordance of life history with genetic and morphological variation suggests that differences between univoltine and bivoltine populations are maintained by selection. Increased electrophoretic variation in the recently colonized range may have arisen by selection on rare variants within the old part of the range.     

Emergence patterns of univoltine and bivoltine Ips typographus (L.) populations and associated natural enemies

Control measures aiming at reducing bark beetle populations and preserving their natural enemies require a sound knowledge on their overwintering and emergence behaviour. These behavioural traits were investigated in univoltine and bivoltine populations of the European spruce bark beetle (Ips typographus [L.], Coleoptera: Scolytinae) and its predators and parasitoids over several consecutive years. In univoltine populations, roughly 50% of the bark beetles left their brood trees in fall together with most parasitoids and some significant predatory flies and beetles. In bivoltine populations, <10% of the second bark beetle generation emerged in fall and the remainder overwintered under the bark of their brood trees. Likewise, most predatory beetles and flies spent wintertime with their prey under the bark, while most parasitic wasps emerged in fall. The spring emergence of bivoltine predatory beetles was found to occur up to 3 weeks earlier than that of I. typographus, while that of the predatory flies and the parasitoids was delayed by up to 1 month. In univoltine populations, the bark beetles emerged several weeks prior to most antagonistic taxa. In the heat year 2003, three I. typographus generations were produced at the lower location, 36% of the third generation emerged in fall, while the proportions of overwintering predators remained largely the same as in previous years. Similar to their host, more parasitoids left their brood trees in fall after warm years. The results show that sanitation felling during winter probably kills most bark beetles in bivoltine populations, but also eliminates many natural enemies. In univoltine populations, sanitation felling might be less detrimental to both I. typographus and natural enemies because a fair fraction of their populations will already have left the trees before cutting. Warmer climates may affect the interactions of bark beetles and natural enemies and thus the impact of control measures.     

Patterns of phenotypic and genetic variation for the plasticity of diapause incidence

Phenotypic plasticity describes an organism's ability to produce multiple phenotypes in direct response to its environmental conditions. Over the past 15 years empiricists have found that this plasticity frequently exhibits geographic variation and often possesses a significant heritable genetic basis. However, few studies have examined both of these aspects of plasticity simultaneously. Here, we examined both the geographic and genetic variations of the plasticity for diapause incidence (the proportion of eggs that enter an arrested state of development capable of surviving over the winter) relative to temperatures and photoperiods associated with long and short season environments across six populations of the striped ground cricket, Allonemobius socius, using a half-sibling split brood quantitative genetic design. We found that plasticity, as measured by the slope of the reaction norm, was greater in the southern-low altitude region (where populations are bivoltine) relative to the southern-high and northern-low altitude regions (where populations are univoltine). However, the heritability of plasticity was only significantly different from zero in univoltine populations that experienced "intermediate" natal season lengths. These patterns suggest that selection may favor the plasticity of diapause incidence in bivoltine regions, but act against plasticity in regions in which populations are univoltine. Furthermore, our data suggest that under "intermediate" natal season length conditions, the interplay between local adaptation and gene flow may keep the plasticity of diapause incidence low (but still significant) while maintaining its genetic variation. As such, this study not only provides a novel observation into the geographic variation of phenotypic plasticity, but also provides much needed groundwork for tests of its adaptive significance.     

Are Polyommatus hispana and Polyommatus slovacus bivoltine Polyommatus coridon (Lepidoptera: Lycaenidae)? The discriminatory value of genetics in taxonomy

Taxonomic confusion exists in several sibling species groups. The Polyommatus coridon species complex (Chalk Hill Blues) serves as a model group of sibling species in which genetic analyses provide suitable means for taxonomic clarification. We studied the allozyme patterns of the two described bivoltine species of this complex, Polyommatus hispana and Polyommatus slovacus, and compared them to the two genetic lineages of the univoltine P. coridon. P. hispana is well distinguished from P. coridon (genetic distance: 0.081), and most probably is a sibling species that has evolved during glacial isolation on the Iberian Peninsula. P. slovacus is genetically indistinguishable from the eastern, Pontic–Mediterranean lineage of P. coridon; therefore we suggest that it represents a local bivoltine population only. Since the spring generation of P. slovacus was much less common than the summer generation and showed less genetic diversity, it is probable that uni-/bivoltinism is a dimorphism affecting only part of the whole population. We suggest that the higher genetic diversity of the second generation may be a consequence of gene flow from adjacent single-brooded populations.     

Larval aestivation and direct development as alternative strategies in the speckled wood butterfly, Pararge aegeria, in Sweden

ABSTRACT.

• 1 Sweden has two disjunct populations of the speckled wood butterfly, Pararge aegeria L. The southern population has two generations per year but the central Swedish population is univoltine. When rearing larvae from central Sweden under normal photoperiodic conditions but at temperatures slightly above the ambient, 42% of the larvae developed directly and produced a second generation of adults the same summer. The egg—larval development time of the directly developing individuals was about 40 days, whereas that of the individuals developing along the univoltine pathway was about 100 days.
• 2 Larvae of the central Swedish population normally aestivate during part of the summer even though abundant food is available. In the closely related Lasiommata petropolitana F., which is the only Swedish satyrid that overwinters in the pupal stage besides P.aegeria, larvae do not aestivate, indicating that there does not seem to be any obligatory association between pupal hibernation and larval aestivation.
• 3 Development rates of aestivating and directly developing P.aegeria are equal up to the third larval instar. During the third and fourth instars, however, the development rate of aestivating individuals is retarded and females also have an additional fifth instar.
• 4 Since the central Swedish P.aegeria have the capacity to develop directly, and the southern Swedish ones have the capacity to aestivate, the evidence indicates that the outcome of the cost/benefit balance of univoltine versus bivoltine development differs between the two areas.

     

ADAPTATION TO SEASONALITY IN A CRICKET: PATTERNS OF PHENOTYPIC AND GENOTYPIC VARIATION IN BODY SIZE AND DIAPAUSE EXPRESSION ALONG A CLINE IN SEASON LENGTH

This paper investigates patterns of phenotypic variation in the striped ground cricket (Allonemobious fasciatus) along a cline in season length and tests the hypothesis that variation in body size and diapause propensity is the result of diversifying selection due to different local conditions. We examined 83 populations and found that A. fasciatus produced a single generation per year (univoltine) north of 37°N latitude and was bivoltine south of 35°30′N. Body size generally increased with increasing season length, with a sudden drop in the region corresponding to the transition from univoltine to bivoltine life cycles, reflecting the division of total season length in two within-bivoltine populations. We reared ten populations in our laboratory and found that much of the interpopulation variance in body size observed in the field could be attributed to genetic differences. Diapause expression also varied significantly among populations and was strongly correlated with season length. The heritability of body size did not differ between populations, but full-sib estimates greatly exceeded parent–offspring estimates suggesting that there are important nonadditive genetic effects. The heritability of diapause expression, determined from analysis of full sibs, varied significantly between populations (0.33 ± 0.10 to 1.31 ± 0.21) with an average of 0.74 ± 0.16. Body size and diapause expression were genetically correlated in transition-zone populations, but not in univoltine or bivoltine populations. Our findings support the suggestion that clinal variation in body size and diapause expression in the striped ground cricket reflect adaptation to season length.     

Climatic adaptation and species status in the lawn ground cricket

Summary The cricket tentatively identified as Pteronemobius taprobanensis shows a stepwise pattern of latitudinal variation in ovipositor length. Abrupt elongation of ovipositor at about 28° N marks the replacement of the subtropical form by the temperate form. The latter maintains almost a constant ovipositor length up to about 35° N within the bivoltine area. Further north in the univoltine area, an ascending cline extends to about 39° N, beyond which no further increase occurs. The ovipositor length adjusted for body size shows a northeastward increase in each of the univoltine and bivoltine areas, though this tendency is less clear in the latter. The optimum length of ovipositor would vary with the relative amounts of gain (due to the protection of eggs in the soil) and loss (due to the metabolic cost and the difficulty to emerge from the soil) in fitness. If so, it may be predicted that a longer ovipositor would be selected for, when the metabolic cost and the risk at hatching are smaller, the environmental pressure in the egg stage is stronger, and the egg stage lasts longer. The last parameter seems to be mainly responsible for the observed association of the type of life cycle with the geographic pattern of variation in ovipositor length.Contribution No. 76 from the Laboratory of Entomology, Hirosaki University     

Life cycles and habitats of wisconsin heptageniidae (ephemeroptera)

Detailed studies were made of the life cycles of Heptageniidae known to occur in Wisconsin. 19 species had univoltine cycles while two and possibly a third had bivoltine cycles. Three univoltine species developed in late spring and early summer while the other univoltine species developed in fall, winter and early spring. For three of the univoltine species, eggs hatched both in fall and the following spring. Diagrams of the life cycles of 15 Wisconsin heptageniids are presented, illustrating the different types of life cycles. Also presented are observation on the food and habitats of the nymphs and flight periods of the adults.     

Comparison on DNA patterns of different ecotypes of European corn borer (Ostrinia nubilalis Hübner) in Hungary

For a molecular genetic study on Hungarian populations of European corn borer L5 stage larvae were collected from 14 places of three different regions of the country (uni- and bivoltine ecotypes). Additionally, the study included larvae from Egypt, too (multivoltine ecotype). Molecular examinations of European corn borer larvae using the study of mitochondrial cytochrome b (cyt b) revealed that by single strand conformation polimorphism (SSCP) the populations found in Hungary represented the same haplotype. Even the Egyptian sample showed no genetic divergence. Some minor deviatons were found in the case of a sample from Székkutas, but that did not prove the genetic divergence of the bivoltine ecotype either, since the other samples of South-East Hungary did not display this kind of genetic variation. On the basis of our investigations it can be said that the univoltine and bivoltine generations, have uniform genetic complements.