Flight Directions in the Migratory Butterfly Pieris brassicae: Results from Semi-natural Experiments

A recently developed experimental set-up has been used to test the constancy of flight direction in the large white butterfly Pieris brassicae, originally collected in Northern Germany. The flight direction, tested with virgin females, depends on the developmental mode. Butterflies from diapause pupae fly north, those from subitan pupae fly south. The constancy of the flight direction has been examined in detail with the spring generation over a period of 14d. The preferred direction does not change with advancing age and is essentially not influenced by temperature or by the sun's azimuth. The flight activity, however, increases with temperature, but this is concealed by an intrinsic response resulting in a diurnal peak of flight activity at 10.00–11.00 hours, long before the maximum temperature of the day is reached. Whether the observed direction can be considered as a parameter of migration behaviour is discussed. It is shown that migration in Pieris brassicae does not end with reproductive maturity but lasts throughout the entire imaginal stage.     

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.     

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.     

Aestivo-hibernal reproductive diapause in Drosophila moriwakii: flight as a controlling factor

Drosophila moriwakii Okada & Kurokawa (Diptera; Drosophilidae) generally has only one generation per year and enters aestivo-hibernal reproductive diapause in Sapporo, northern Japan, but a small fraction of the population produced a second generation in summer at a place where breeding resources were abundant. In this species, diapause seems to be controlled by flight activity. When flies were cultured in cages in which they were able to fly freely, they entered diapause irrespective of photoperiod, but they did not do so at long daylengths when cultured in small vials in which they were prevented from flying. Furthermore, flies with wings removed did not enter diapause at long daylengths even if they were cultured in the cages.     

A new experimental approach to investigate migration in Pieris brassicae L.

Abstract.

• 1 It is possible to correlate distinct sequences of flight behaviour in caged females of the migrant butterfly Pieris brassicae with the main flight direction of released individuals of the same stock. The attempts to escape out of the cage point to the same mean direction as the flights of the released butterflies. This provides an experimental tool to investigate some unsolved questions concerning migration behaviour.
• 2 This study examines the constancy of the flight direction of a group of butterflies, the specificity of the direction in two populations of different geographical origin, and the frequently reported change in flight direction of subsequent generations.
• 3 The results indicate that the main flight direction of a tested group is very constant during 2 days of observation. Different directions have been detected in populations originating from Northern Germany and Southern France, respectively. The preferred directions are discussed as an adaptation to the geographical circumstances. Subsequent generations of the population from Northern Germany also show different directions. The spring generation flies to the north, the autumn generation in the opposite direction. This result corresponds to field observations that in P.brassicae a return flight occurs in subsequent generations of a year.

     

Seasonal butterfly design: morphological plasticity among three developmental pathways relative to sex,flight and thermoregulation

The temperate‐zone butterfly Pararge aegeria can use three developmental pathways corresponding to different seasonal cohorts: (1) development with a pupal winter diapause resulting in early spring adults; (2) development with a larval winter diapause resulting in late‐spring adults and (3) direct development resulting in summer or second generation adults. In order to test adaptive predictions, we compared variation in flight‐ and thermoregulation‐related morphology among adult males and females from the three pathways using both field data (i.e. wild‐caught butterflies) and experimental breeding data (i.e. reared under different photoperiod regimes). Morphological patterns among the pathways were largely similar in the field and rearing data. Seasonal patterns differed between the sexes for most traits, including (relative) size measures and wing colour. Our results suggest sex‐related, adaptive seasonal plasticity for morphological traits related to flight behaviour in a multivoltine insect.     

Ground crickets singing in volcanic warm “islets” in snowy winter: Their seasonal life cycles,photoperiodic responses and origin

The ground cricket Dianemobius nigrofasciatus overwinters as an egg in Japan, being univoltine in Hokkaido and northern Honshu and bivoltine farther south. In Hokkaido, however, this cricket is heard singing in winter in several fumarolic fields covered with moss and grasses locally known as “bokke”. In such warm “islets” the adult density was high in early summer and again in autumn, indicating that the cricket is bivoltine in contrast to the univoltine life cycle outside the bokke habitats in Hokkaido. Eggs laid by females collected at regular intervals from a bokke habitat showed a clear seasonal cycle of diapause incidence. At 26°C, the bokke strains produced non‐diapause eggs under long days and diapause eggs under short days as in the southern bivoltine populations, although the critical day‐length was longer than in the south. Several strains derived from non‐bokke habitats in Hokkaido and northern Honshu produced high percentages of diapause eggs under long days as well as short days as expected for the univoltine life cycle. Winter adults singing in bokke habitats could be either survivors of the autumn generation or individuals derived from eggs laid in autumn and then matured in response to the high soil temperature. In the laboratory, the proportion of egg diapause in short days was decreased by selection only for several generations. Phylogenetic trees of bokke and non‐bokke populations, based on both the nucleotide sequence of the mitochondrial COI gene and four allozyme loci, suggest that bokke populations have not been isolated from non‐bokke populations for an evolutionarily significant time.     

Geographic variation in diapause induction and termination of the cotton bollworm, Helicoverpa armigera Hübner (Lepidoptera: Noctuidae)

Overwintering diapause in Helicoverpa armigera, a multivoltine species, is controlled by response to photoperiod and temperature. Photoperiodic responses from 5 different geographical populations showed that the variation in critical photoperiod for diapause induction was positively related to the latitudinal origin of the populations at 20, 22 and 25 °C. Diapause response to photoperiod and temperature was quite different between northern and southern populations, being highly sensitive to photoperiod in northern populations and temperature dependence in southern populations. Diapause pupae from southern population showed a significantly shorter diapause duration than from northern-most populations when they were cultured at 20, 22, 25, 28 and 31 °C; by contrast, overwintering pupae from southern populations emerged significantly later than from northern populations when they were maintained in natural conditions, showing a clinal latitudinal variation in diapause termination. Diapause-inducing temperature had a significant effect on diapause duration, but with a significant difference between southern and northern populations. The higher rearing temperature of 22 °C evoked a more intense diapause than did 20 °C in northern populations; but a less intense diapause in southern population. Cold exposure (chilling) is not necessary to break the pupal diapause. The higher the temperature, the quicker the diapause terminated. Response of diapause termination to chilling showed that northern populations were more sensitive to chilling than southern population.     

The role of gene flow asymmetry along an environmental gradient in constraining local adaptation and range expansion

Theoretically, asymmetric gene flow along an environmental gradient can limit species range expansion by keeping peripheral populations from locally adapting. However, few empirical studies have examined this potentially fundamental evolutionary mechanism. We address this possibility in the cricket Allonemobius socius, which exist along a season‐length gradient where the probability of producing a single generation per year (univoltinism) increases with latitude. As the probability of univoltinism increases northwards, populations are expected to hedge their bets by producing a greater proportion of diapause eggs when exposed to a mild diapause cue. However, gene flow from southern populations may disrupt local adaptation in the north by reducing the proportion of diapause eggs (expected to be 100% in pure univoltine environments). This may limit range expansion along the northern periphery where A. socius compete with A. fasciatus, a sister species that exhibits an invariant diapause‐only egg‐laying strategy. To assess the potential for range limitation, we examined diapause incidence (the proportion of diapause eggs produced under diapause conditions), gene flow symmetry and population structure across nine A. socius populations. We found that gene flow was asymmetric and biased northwards towards the periphery. Furthermore, peripheral populations that inhabited pure univoltine environments produced numerous nondiapause eggs (a southern, bivoltine diapause phenotype), which we assume to be a suboptimal phenotype. These patterns suggest that asymmetric gene flow along the gradient constrains adaptation in peripheral populations, potentially constraining species range expansion.     

中华稻蝗长沙种群的生活史及其卵滞育的进化意义

中华稻蝗Oxya chinensis为重要的水稻害虫,在我国除青海、西藏、新疆、内蒙古等未见报道外,南起海南北至东北均有分布,在许多的分布区域1年发生1代。为探索中华稻蝗长沙种群的生活史及其季节适应特征,通过野外和实验室的研究,调查了其发生代数、孵化率和卵滞育率的季节变化及越冬卵的存活率。结果显示,中华稻蝗长沙种群为1年2代和1年1代混合发生:第1代卵产卵后大部分孵化为若虫而1年完成2代,但亦有19.4% -4.1%的卵不孵化而1年只能完成1世代。第1代成虫于6月上旬至8月上旬羽化,6月下旬至8月中旬产卵;第2代若虫于7月初开始孵化,9-10月羽化为成虫,10月上旬至11月下旬产卵。在室外自然条件下,中华稻蝗长沙种群6-8月(第1代)和10-11月(第2代)所产卵块均为部分滞育,滞育率为30%左右,皆无显著差异。然而,其卵滞育率在12月以后显著降低,仅为6.6%或以下,卵滞育快速地得以解除。因此,包括非滞育卵和滞育解除卵,中华稻蝗长沙种群的越冬卵皆以非滞育状态度过其后的寒冷季节。即使是遭遇长江流域2007年末至 2008年初异常寒冷的冬季,在长沙地区越冬后其卵的存活率亦在98%以上。非滞育状态的中华稻蝗长沙种群越冬卵完全能安全地越冬,其滞育的发生并非是为了提高其耐寒性而安全度过不适环境。并探讨了中华稻蝗长沙种群卵滞育的进化意义。     

Heterodynamic processes in Coccinella septempunctata L. (Coccinellidae: Coleoptera): a mini review

Among ladybirds, diapause has evidently been most studied in Coccinella septempunctata L. adults. Coccinella septempunctata (C‐7), the seven spot, has attracted attention both in Europe and the USA for being heterogeneous as to the induction of diapause. A polyphenic character of diapause is a prominent feature in C. septempunctata and this phenomenon generally and often generates voltinism heterogeneity within populations. A greater part of the C‐7 population shows an obligatory univoltine cycle, whereas a smaller proportion is facultatively polyvoltine. Coccinella s. bruckii in Japan is bivoltine, in which the first generation of adults aestivate while the second generation hibernates. This paper reviews this heterodynamic cycle and its importance in the life history of C‐7, as reported from different regions of the world. Heterogeneous voltinism and diapause smooth the progress of a portion of C‐7 populations to take benefit of aphids well into late summer and autumn. Understanding such variation and plasticity in the life history of this species can help in organizing proper biological control attempts using it as a biocontrol agent.     

Geographical variability in ecophysiological traits controlling dormancy in Chrysopa oculata (Neuroptera: Chrysopidae)

Dormancy in Chrysopa oculata, a multivoltine species, is controlled by reponses to photoperiod and temperature. Variation in both the critical photoperiod for diapause induction and the duration of diapause is positively related to the latitudianl origin of the population. The incidence of photoperiodically induced diapause also varies geographically; at 24°C the southern-most population is more variable for diapause induction than the northern populations. By contrast, there is little interpopulation variation in thermally determined rates of post-diapause development. The results suggest that the lower threshold for development and the thermal requirements above the threshold may be functionally correlated.     

Life‐history traits related to diapause in univoltine and bivoltine populations of Ypthima multistriata (Lepidoptera: Satyridae) inhabiting similar latitudes

In most temperate insects, diapause strategies and voltinism generally exhibit latitudinal clines, supporting the concept that they represent adaptations to climate. In contrast, in the satyrine butterfly Ypthima multistriata Butler, local populations with different voltinism patterns are geographically intermingled, suggesting that life‐history traits related to diapause may differ even between geographically and phylogenetically close populations. In this study, we experimentally examined the critical photoperiod for diapause induction and the larval developmental period in two univoltine and two bivoltine populations of Y. multistriata, all of which inhabit virtually the same latitude (34.652–34.750°N). We found that the critical photoperiod for diapause induction was longer in the univoltine populations than in the bivoltine populations. Moreover, the larval period under the long day length treatment was different among populations in both sexes, although significant differences were also detected between populations with the same voltinism. These results indicate that in Y. multistriata, life‐history traits related to diapause can not be attributed merely to climatic conditions such as temperature or day length, which depend largely on latitude. Therefore, we suggest that biotic elements, such as leaf toughness, as well as abiotic elements should be taken into account in attempts to explain the enigmatic pattern of geographic variation in the diapause strategies of Y. multistriata.     

稻水象甲(Lissorhoptrus oryzophilus Kuschel)的卵子发生-飞行共轭

１９９４～１９９６年对双季稻区稻水象甲发生消长和迁飞动态的系统研究表明,稻水象甲的飞行肌和卵巢发育随生活史的季节性变化出现兴衰交替,从而迁入迁出稻田和越冬场所,表现出典型的卵子发生－飞行共轭：春季越冬代成虫从越冬场所迁入早稻田后飞行肌消解而卵巢发育,繁殖形成一代致害种群。夏季一代成虫生殖滞育,飞行肌和脂肪体发达,绝大部分个体迁出早稻田行夏蛰并越冬;少量落入秧田者和早稻收割时散落田内而晚稻插秧时尚未迁离的个体卵巢恢复发育,飞行肌消解而构成二代虫源。秋季二代成虫羽化后生殖滞育,迁飞上山越冬;１０月中旬后羽化的个体卵巢和飞行肌均不再发育而滞留田内外越冬。取食高质量食料时,一代夏蛰成虫飞行肌再生,新羽一代成虫则首先发育飞行肌,而后解除生殖滞育。     

Rapid seasonal adaptation of an alien bruchid after introduction: geographic variation in life cycle synchronization and critical photoperiod for diapause induction

Whether alien insects that are introduced into temperate regions adapt to seasonally changing environmental conditions is an important question in evolutionary biology. If rapid evolution has occurred in a non‐native environment, a latitudinal cline in critical photoperiod for diapause induction (i.e., the photoperiod at which half of the individuals enter diapause) and in life cycle synchronization with host plant phenology should be evident among locations. The alien bruchid Acanthoscelides pallidipennis (Motschulsky) (Coleoptera: Bruchidae) is native to North America and introduced into Japan with the host plant Amorpha fruticosa L. (Fabaceae) in the late 1940s. To examine whether seasonal adaptation has occurred in A. pallidipennis, we conducted a laboratory experiment and phenological observations using three latitudinally different populations. We bred F1 eggs at 22 °C and five photoperiodic regimens – L:D = 10:14, 13:11, 14:10, 15:9, or 16:8 hours – and examined whether diapause was induced. The estimated critical photoperiod for diapause induction was longest in the most northern population and shortest in the most southern population. Life cycle was found to be synchronized with host phenology in each location. Also voltinism varied geographically, from univoltine in the northern population to bivoltine in the southern populations. These results showed that A. pallidipennis rapidly adapted to seasonal environmental conditions in Japan after its introduction.     

Survivability and post‐diapause fitness in a scolytid beetle as a function of overwintering developmental stage and the implications for population dynamics

1. It has recently been suggested that expected increases in temperature might lead to an additional generation per season in bark beetles. Thus, populations would grow more rapidly. However, an additional but not fully developed generation could lead to high winter mortality in the pre‐imaginal stages. 2. Winter survivability and post‐diapause fitness as a function of the overwintering developmental stage of the spruce bark beetle Ips typographus L. were studied along an altitudinal cline. Fitness was tested in terms of weight, lipid content and flight capacity. 3. Mortality was significantly lower and emergence per brood system significantly higher for fully developed adults that entered overwintering than for larvae, pupae or callow adults. 4. Post‐diapause fitness in terms of dry weight and flight capacity was significantly higher in individuals that completed development before winter, and lipid contents also showed a trend for being higher in those individuals. 5. In conclusion, in a scenario where effective temperature sums are not adequate for the complete development of an additional generation, models may overestimate population growth by neglecting increased mortality and reduced post‐diapause fitness. The results highlight the importance of considering life‐history traits and indirect effects in addition to abiotic factors such as temperature when modelling population dynamics.     

Geographic variation in critical photoperiod for diapause induction and its temperature dependence in Hyphantria cunea Drury (Lepidoptera: Arctiidae)

The fall webworm, Hyphantria cunea Drury (Lepidoptera: Arctiidae), was introduced from North America to Japan half a century ago. The critical photoperiod for diapause induction and its temperature dependence, as defined by the difference in the critical photoperiod between 20 and 25°C, were investigated in order to understand the mechanisms behind a shift from bi- to trivoltine life cycles. The critical photoperiod for diapause induction was shorter in the southern trivoltine populations than in the northern bivoltine populations, and this was more marked at 25°C than at 20°C. Although the critical photoperiod showed a positive correlation with the original latitude, the correlation was relatively low at both temperatures. Conversely, temperature dependence of the critical photoperiod for diapause induction correlated negatively with the original latitude. The trivoltine populations showed greater temperature sensitivity than the bivoltine populations. These results suggest that an increase in temperature sensitivity of the diapause response to photoperiods was involved in the shift to a trivoltine life cycle. The crossing experiments suggested that the photoperiodic control of diapause induction and its temperature dependence are under polygenic control without sex-linkage. Received: 29 October 1996 / Accepted: 26 February 1997     

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.     

Effects of climatic change on the phenology of butterflies in the northwest Mediterranean Basin

Phenological changes in response to climatic warming have been detected across a wide range of organisms. Butterflies stand out as one of the most popular groups of indicators of climatic change, given that, firstly, they are poikilothermic and, secondly, have been the subject of thorough monitoring programmes in several countries for a number of decades. Here we provide for the first time strong evidence of phenological change as a consequence of recent climatic warming in butterflies at a Spanish site in the northwest Mediterranean Basin. By means of the widely used Butterfly Monitoring Scheme methodology, three different phenological parameters were analysed for the most common species to test for trends over time and relationships with temperature and precipitation. Between 1988 and 2002, there was a tendency for earlier first appearance dates in all 17 butterfly species tested, and significant advances in mean flight dates in 8 out of 19 species. On the other hand, the shape of the curve of adult emergence did not show any regular pattern. These changes paralleled an increase of 1–1.5°C in mean February, March and June temperatures. Likewise, a correlation analysis indicated the strong negative effect of spring temperature on phenological parameters (i.e. higher temperatures tended to produce phenological advances), and the opposite effect of precipitation in certain months. In addition, there was some evidence to indicate that phenological responses may differ between taxonomic lineages or species with similar diets. We discuss the consequences that these changes may have on species' population abundances, especially given the expected increase in aridity in the Mediterranean Basin caused by current climatic warming. We predict that varying degrees of phenological flexibility may account for differences in species' responses and, for multivoltine species, predict strong selection favouring local seasonal adaptations such as diapause phenomena or migratory behaviour.     

Latitudinal clines in the timing and temperature-sensitivity of photoperiodic reproductive diapause in Drosophila montana

Reproductive diapause is a primary mechanism used by arthropods to synchronize their life cycle with seasonal changes in temperate regions. Our study species, Drosophila montana, represents the northern insect species where flies enter reproductive diapause under short day conditions and where the precise timing of diapause is crucial for both survival and offspring production. We have studied clinal variation in the critical day length for female diapause induction (CDL) and their overall susceptibility to enter diapause (diapause incidence), as well as the temperature sensitivity of these traits. The study was performed using multiple strains from four latitudinal clines of the species – short clines in Finland and Alaska and long clines in the Rocky Mountains and the western coast of North America – and from one population in Kamchatka, Russia. CDL showed strong latitudinal clines on both continents, decreasing by one hour per five degrees decline in latitude, on average. CDL also decreased in all populations along with an increase in fly rearing temperature postponing the diapause to later calendar time, the effects of temperature being stronger in southern than in northern population. Female diapause incidence was close to 100% under short day/low temperature conditions in all populations, but decreased below 50% even under short days in 19°C in the southern North American western coast populations and in 22°C in most populations. Comparing a diversity of climatic data for the studied populations showed that while CDL is under a tight photoperiodic regulation linked with latitude, its length depends also on climatic factors determining the growing season length. Overall, the study deepens our understanding of how spatial and environmental parameters affect the seasonal timing of an important biological event, reproductive diapause and helps to estimate the evolutionary potential of insect populations to survive in changing climatic conditions.