Poly modal emergence of the tiger swallowtail, Papilio glaucus (Lepidoptera: Papilionidae): source of a false second generation in central New York State

Abstract. 1. The tiger swallowtail butterfly, Papilio glaucus (L.) (Lepidoptera: Papilionidae), is commonly assumed to have a bivoltine life cycle in central New York State and other parts of the northeastern United States. The seasonal abundance of adults in Tompkins County, New York, shows a bimodal pattern in some years, and a skewed unimodal pattern in others. We studied the incidence of diapause, the potential for successful development of two generations in the field, and the dates of adult emergence from overwintering pupae to determine whether these seasonal patterns are the result of bivoltinism.
2. Insects from Tompkins County, reared as larvae under 16L:8D photoperiod, uniformly enter pupal diapause.
3. Comparison of the heat units required for successful development of two generations with the heat units available in the field during 11 years shows that emergence of non-diapausing individuals would occur too late in the summer to contribute to the second peak of butterfly abundance. Offspring of non-diapausing individuals could only rarely complete development before the end of the growth season.
4. Diapausing offspring of butterflies collected in early June and late July differed sufficiently in dates of emergence to account for the field pattern of seasonal abundance.
5. Electrophoretic studies revealed no evidence of genetic isolation between early and late emerging butterflies.
6. These results suggest that polymodal emergence of diapausing insects can masquerade as a bivoltine life cycle.     

How climate change affects the occurrence of a second generation in the univoltine Pyrrhocoris apterus (Heteroptera: Pyrrhocoridae)

1. Understanding the conditions that allow for the occurrence of an additional generation in populations that are usually univoltine is important under the present climate warming. In temperate areas, a second generation is enabled through the emergence of a time window that opens when first-generation individuals are ready to reproduce and closes when second-generation individuals cannot complete development before the onset of winter. 2. The conditions that limit the width of this window were studied in Pyrrhocoris apterus (Heteroptera: Pyrrhocoridae), a ground-inhabiting heteropteran overwintering in facultative adult diapause, whose populations in Central Europe have typically been univoltine until the 1980s. 3. The frequency of females of the first generation that started to lay eggs decreased from 70% in June to zero in early August, but oviposition of these females continued until the end of August. Using thermal constants for egg–adult development and temperature data, this study found that the development of most second-generation individuals could only be completed before the start of winter if hastened through behavioural thermoregulation. 4. Consequences of temperature increase on the width of the thermal window were calculated. Increasing temperature causes the time window to open earlier and close later by accelerating maturation of first-generation females and improving conditions for maturing of the second-generation individuals in late summer and autumn. 5. Climate warming will create conditions that facilitate the occurrence of a second generation in a year in typically univoltine populations of this species.     

Timing and duration of the life cycle of Asian longhorn beetle Anoplophora glabripennis (Coleoptera: Cerambycidae) in southern England

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Temperature‐size responses match latitudinal‐size clines in arthropods,revealing critical differences between aquatic and terrestrial species

Two major intraspecific patterns of adult size variation are plastic temperature‐size (T‐S) responses and latitude‐size (L‐S) clines. Yet, the degree to which these co‐vary and share explanatory mechanisms has not been systematically evaluated. We present the largest quantitative comparison of these gradients to date, and find that their direction and magnitude co‐vary among 12 arthropod orders (r² = 0.72). Body size in aquatic species generally reduces with both warming and decreasing latitude, whereas terrestrial species have much reduced and even opposite gradients. These patterns support the prediction that oxygen limitation is a major controlling factor in water, but not in air. Furthermore, voltinism explains much of the variation in T‐S and L‐S patterns in terrestrial but not aquatic species. While body size decreases with warming and with decreasing latitude in multivoltine terrestrial arthropods, size increases on average in univoltine species, consistent with predictions from size vs. season‐length trade‐offs.     

The effect of temperature on the development and life cycle regulation of the pine weevil Hylobius abietis and the potential impacts of climate change

• 1 The pine weevil Hylobius abietis is widely distributed in the Palaearctic region where it is a major pest. Although predominantly semi‐voltine, with a 2‐year life cycle, the generation time across its range can vary from 1 to 4 years. The duration of the life cycle and the seasonal timing of weevil activity affect the economic impact and management of this pest, all of which are likely to change in a warming climate.
• 2 To determine the effect of temperature and tree species on weevil growth and development, laboratory experiments were performed with eggs, larvae, prepupae, pupae and adults, using, as appropriate, the host species Scots pine Pinus sylvestris L. and Sitka spruce Picea sitchensis (Bong.) Carr. under constant or alternating temperatures.
• 3 The development rate was linearly related to temperature, with developmental thresholds for eggs, larvae and pupae of 8, 4.5 and 7.3 °C, respectively. Day‐degrees were estimated for each life stage. Larval development was affected by tree species, being slower on Sitka spruce than on Scots pine, and was faster under alternating than constant temperatures.
• 4 The development time for prepupae was highly variable, with an apparent facultative prepupal diapause initiated by temperature. The temperature range 20–17.5 °C marked the transition between median prepupal development times of approximately 25 and 90 days. The prepupal stage may serve to minimize the risk of overwintering mortality in the pupal stage and help to synchronize the life cycle.
• 5 Larval and adult mass was positively related to developmental temperature, demonstrating an inverse temperature size rule, and weevils were heavier when developing on Scots pine than Sitka spruce. Development in alternating temperatures reduced weevil mass on Scots pine. The influence of temperature on weevil mass is likely to have a positive effect on fecundity and overwintering survival. The effects of climate change on development, voltinism and weevil mass are discussed.

     

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.     

Countergradient vs. cogradient variation in growth and diapause in a lichen‐feeding moth,Eilema depressum (Lepidoptera: Arctiidae)

Separating genetic and environmental causes of the latitudinal differences among populations is crucial when evaluating the potential for microevolutionary responses to the changing environment. We studied among‐population and environmental components of variation in several life‐history traits of a lichen‐feeding moth Eilema depressum when offspring of replicate Swiss and Finnish females were reared in a common‐garden factorial experiment. A partial second generation was produced only among Swiss larvae, more likely so at higher temperature regime and higher host quality, and more frequently among the offspring of particular females. Growth rates of larvae that chose the diapause development were higher in northern individuals. Our results thus reveal adaptive differences between latitudinal populations in studied life‐history traits, allowing to expect rapid adaptation of the species to further environmental changes. In contrast, invariable responses of the growth rates of the larvae to temperature and host quality support the idea that some basic parameters of insect growth show a high degree of evolutionary conservatism.     

Variability in the life history of the water strider Gerris lacustris (Heteroptera: Gerridae) across small spatial scales

Abstract.  1. Variability in life-history traits of the common pond skater Gerris lacustris (L.) (Heteroptera: Gerridae) was investigated on a small geographical scale in Bavaria, Germany (≈ 49 °N). In an area of approximately 3.5 × 3.0 km, voltinism and wing length data of G. lacustris were recorded from 65 ponds located either in an open field or surrounded by forest.
2. Life history differed significantly between populations in the two habitat types. Gerris lacustris showed a bivoltine life-cycle in most of the field ponds with a high proportion of long-winged individuals in the diapause generation and a large increase of the short-winged morph in the midsummer generation. In contrast, almost all populations inhabiting forest ponds were univoltine. Only a few forest populations produced any second generation larvae. The long-winged morph dominated during the whole season in forest ponds.
3. The appearance of first-instar larvae was delayed by 3–4 weeks in forest ponds compared with field ponds. Presumably, low temperatures led to this reduction in the developmental rate of eggs and larvae. Consequently, 'field larvae' of the first generation developed under increasing daylight conditions (triggering direct reproduction and brachyptery) but 'forest larvae' developed under a shortening photoperiod (triggering diapause and macroptery).
4. Offspring raised from 'forest parents' in the laboratory showed the same variability in life history when reared in a cool (16.9 ± 2.4 °C) or warm (23.1 ± 1.8 °C) temperature regime.
5. Given the small spatial scale of the study, it is concluded that the variability in the life history of G. lacustris has a strong phenotypic component.     

Control of diapause induction by a change in photoperiod in Melanoplus sanguinipes

The effect of photoperiod on diapause induction, developmental time and body size was examined in Melanoplus sanguinipes, the lesser migratory grasshopper. Contrary to what is found in most insects, facultative diapause-egg production is found to be controlled by changing rather than constant photoperiods. In addition, developmental time is shown to be faster under short-day photoperiods than long-day photoperiods. And finally, body size is larger under long-day photoperiods and smaller under short-day photoperiods. Implications of these results for the regulation of the seasonal life cycle of M. sanguinipes in the field are discussed.     

Photoperiod- and temperature-dependent induction of larval diapause in a multivoltine bruchid, Bruchidius dorsalis

The wild bruchid beetle, Bruchidius dorsalis Fahraeus (Coleoptera: Bruchidae), has a multivoltine life cycle and overwinters in several developmental stages in the middle part of Japan. We investigated the incidence of diapause under different conditions of photoperiod (from L8:D16 to L16:D8) and temperature (at 20 °C and 24 °C). Our experiments revealed the following results: (1) B. dorsalis entered diapause at the final (late fourth) instar larva under short photoperiods, (2) the larval diapause incidence was dependent on temperature (critical photoperiods were 12.5 h at 20 °C and 12 h at 24 °C), (3) some individuals did not enter diapause under short-photoperiod conditions at 24 °C, and (4) the sensitive stages to the photoperiod were from the late egg stage to the early first instar larva. Based on these results, we discuss not only the evolution of a complex overwintering strategy inB. dorsalis but also the domestication process of stored-bean pests.