Feeding insects with a phytopathogenic fungus influences their diapause and population dynamics

Abstract.  1. A facultative mutualistic relationship between a herbivorous moth, Lobesia botrana , and a phytopathogenic fungus, Botrytis cinerea , both damaging the same plant, the vine, has been demonstrated recently. Laboratory and field studies were carried out to determine the influence of the presence of the fungus in food on larvae during the pre- and post-diapause periods and on the global fitness of the insect.
2. During the pre-diapause period, larvae fed on fungus exhibited a higher survival rate and a faster larval development than insects reared without fungus.
3. After the wintering diapause, insects reared with fungus displayed a better synchronisation of adult emergence and an increased fecundity compared with control insects.
4. These results indicate that food containing fungus provides an improved diet, permitting the herbivorous insect to obtain nutrients required to optimise their life-history traits.
5. In addition, the results emphasise the importance of the nutritional quality of larval diet, improved by addition of the fungus, on diapause termination and on the demography of the insect before and after the wintering diapause.     

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.     

Seasonal susceptibility of 'Bartlett' pears to codling moth (Lepidoptera: Tortricidae) infestation and notes on diapause induction

Seasonal susceptibility of 'Bartlett' pear, Pyrus communis L., to codling moth, Cydia pomonella (L.), infestation, successful completion of larval development after infestation, and the induction of C. pomonella diapause was studied from 1992 through 1995. The seasonal variation in C. pomonella infestation and larval survival were effected by changes in fruit maturity. In late May through mid-June, pears were hard and were not as successfully infested by C. pomonella and produced less larvae compared with fruit later in the season. In late June to mid-July, pears became more suitable for infestation and a greater percentage of the larvae completed their development. In late July through mid-August, pears were susceptible to infestation, but the larvae were less likely to successfully complete development than in the late June to mid-July period due to pear tissue breakdown. From mid-August through September, pears are unsuitable for infestation, and few larvae were produced. When fruit were infested with neonate larvae in late May and mature larvae emerged from the fruit in July, a low percentage of the larvae entered diapause. However, when fruit were infested with neonate larvae in early July and mature larvae emerged from the fruit in early August, the majority of the larvae entered diapause. When fruit were infested with neonate larvae in late July through September and mature larvae emerged from the fruit after mid-August, nearly all C. pomonella larvae had entered diapause.     

Timing of diapause initiation, metabolic changes and overwintering survival of the spruce budworm, Choristoneura fumiferana

1. Four groups of spruce budworm larvae, Choristoneura fumiferana , of the same physiological stage (at the beginning of diapause) were exposed to natural temperatures, starting in July, August, September and October. Post-diapause emergence and certain metabolites were monitored throughout the overwintering period.
2. Larval exposure to high temperatures for long periods before winter had a significant effect on temperature requirements for subsequent diapause development and apparently caused a certain amount of mortality, especially for those larvae that entered diapause earlier in the season.
3. Substantial loss of lipid and glycogen reserves was related to intensity of high temperature exposure during the pre-winter period, which in turn was associated with the timing of diapause initiation.
4. Patterns of glycerol accumulation during the winter were also influenced by temperature conditions in the early phase of diapause development.
5. Among several possible mortality factors, an extended cool period in spring was suggested as a potential cause of overwintering mortality, which was closely associated with a deficiency in energy reserves caused by high temperature exposure before winter.
6. These results stress the significant impact of pre-winter conditions on the overwintering process and highlight the importance of an ecophysiological approach to insect overwintering biology.     

The causes of seasonal changes in numbers of the yellow dung fly, Scathophaga stercoraria (Diptera: Scathophagidae)

ABSTRACT. 1. Counts of adult Scathophaga stercoraria (L.) on cow pats were made in Houghall, County Durham, in 1964 and 1965.
2. A spring peak of numbers was due to adults (overwintered mainly as pupae or larvae) maturing and going to dung to breed. Numbers then dropped, rising to one or more peaks in late June—early July. In 1964 there was then a summer drop in numbers until late September. In 1965 high numbers persisted in summer associated with cooler, wetter weather. Autumn peaks in both years persisted until severe frosts or snow.
3. Mature adults, developed from eggs laid during the spring peak, form the first generation when breeding in mid-late June. No clear generations can be identified after this, due to eggs being laid daily (females have successive gonotrophic cycles). Changes in adult numbers breeding reflects survival of eggs and newly-hatched larvae 5–6 weeks earlier, and lower survival rates of adults in mid-summer compared with spring and autumn.
4. Adult Scathophaga numbers in vegetation rose as numbers on dung dropped. Females dissected to count ovariole tunica dilatations showed that most flies in vegetation were immature, with some parous flies hunting insects to develop the next batch of eggs.
5. Females on dung were dissected and found to range from immature to seven-parous. Those gravid for the first time were grossly under-represented, possibly due to wider dispersal.
6. It is suggested that seasonal changes in this r-strategist cannot be explained simply in terms of generations nor by the occurrence of adult diapause.     

Temperature and photoperiodic control of diapase induction in the ant <Emphasis Type="Italic">Lepisiota semenovi</Emphasis> (Hymenoptera,formicidae) from Turkmenistan

The ants L. semenovi has been found to belong to species with endogenous-heterodynamic seasonal life cycles with the obligate diapause induced predominantly by factors internal for a colony, whereas external ecological factors (photoperiods and temperature) produce merely modifying effects by accelerating or delaying the diapause onset. The photoperiodic and temperature regulation of diapause induction in larvae and queens is shown. Under effect of short days and low temperature the periods of larval pupation and queen oviposition in a colony are shortened markedly, i.e., the diapause of larvae and queens occurs earlier. The daily rhythms of temperature 15/25°C and particularly 20/30°C as compared with constant temperatures 20 and 25°C that correspond to the mean circadian temperatures of the thermorhythm, inhibit manifestations of the short day effects by stimulating the non-diapause development and increasing duration of the seasonal development cycle of ant colonies. The L. semenovi photoperiodic reaction is quantitative, as development and pupation of larvae and egg-laying of queens cease sooner or later under both the short and the long days, but in the latter case significantly later. Thus L. semenovi is one more example among very rare ant species that are revealed to have the photoperiodic regulation of the colony development seasonal cycle.     

Repeated cycles of chilling and warming effectively terminate prolonged larval diapause in the chestnut weevil, Curculio sikkimensis

Curculio sikkimensis (Coleoptera: Curculionidae) requires one or more years to complete its life cycle, owing to prolonged larval diapause. To compare the effects of temperature cycles and total periods of chilling on the termination of prolonged diapause, larvae were subjected to different chilling (5 degrees C) and warming (20 degrees C) cycles ranging from 30 to 720 days, and all cycles were repeated until the sum of chilling and warming periods reached 720 days. The prolonged diapause of C. sikkimensis was more effectively terminated by repeated cycles of chilling and warming than by prolonging the continuous chilling period. However, extremely short temperature cycles were not highly effective in enhancing diapause termination, even when such cycles were repeated many times. To examine the role of warming periods on diapause termination, diapause larvae were subjected to a sequence of chilling (120 days at 5 degrees C) and warming (240 days at 20 degrees C) with a warming period (0-120 days at 20 degrees C) inserted in the chilling period. Diapause larvae that were not reactivated in the first chilling period required exposure to a certain period of warming before they were able to complete diapause development in the subsequent chilling. Thus, C. sikkimensis appears to spread its reactivation times over several years in response to seasonal temperature cycles.     

Seasonal changes in the reproductive and life-history traits of Chrysomela populi L. (Coleoptera: Chrysomelidae)

Seasonal changes in the reproductive and life‐history traits of Chrysomela populi L. were investigated in Shobara (34°52′N, 133°01′E), Japan, in 2004 and 2005. Larvae were hatched and reared under natural photoperiod and temperature in 8 different periods between late April and early September in 2004. The incidence of adult diapause increased with progression of the season in 2004. The critical timing of diapause induction, as defined by 50% of females entering diapause, was estimated to occur between late July and early August. No effects of season on the survival rate from hatching to adult emergence, percentage ovipositing females and percentage females that deposited viable eggs were clearly detected. Adult weight at emergence fluctuated seasonally, which was probably caused by air temperature. No effect of season on the oviposition period was observed among females that averted diapause. Females that entered diapause in 2004 started oviposition from the first day of May in 2005. Reproductive output was significantly greater in diapause females than in non‐diapause females. This increase in the reproductive output of diapause females was caused by elongation of the oviposition period and consequently by enlargement of the number of egg clutches deposited. These results suggest that the seasonal timing of diapause induction in females of C. populi would be affected by various risks and benefits to reproduction and survival.     

Photoperiodic induction of prepupal diapause and its role in synchronization with host phenology in the hibiscus caterpillar,Xanthodes transversa

The hibiscus caterpillar, Xanthodes transversa (Lepidoptera: Noctuidae), is a multivoltine insect that is an important pest of Malvaceae plants such as the okra, Abelmoschus esculentus, and the common rose mallow, Hibiscus mutabilis, in Japan. In the present study, the effects of photoperiod and temperature on the induction of prepupal diapause and the adaptive significance of this diapause were examined in a local population of X. transversa in Miyazaki, Kyushu, southwestern Japan. Larvae showed a long‐day photoperiodic response for controlling the induction of prepupal diapause with a critical day length between 13 and 14 h at 20 to 25°C. Under long‐day conditions larvae rapidly pupated from the sixth instar, but under short‐day conditions they entered diapause in the prepupal stage. Diapause occurrence in the field increased in late September, which was consistent with the laboratory results in terms of the photoperiodic response. Non‐diapause development after this time is maladaptive because most larvae of the next generation could not reach the critical stage (prepupae) before winter and died during early winter in outdoor experiments. Larvae suffered from a high rate of mortality when fed leaves collected late in autumn. The photoperiodic response for controlling the induction of diapause in this insect may play an important role in synchronizing the life cycle with the seasonal changes in food and temperature conditions.     

Photoperiodic control of larval development in the semivoltine cockroachPeriplaneta japonica (Blattidae: Dictyoptera)

Periplaneta japonica is semivoltine, entering early diapause in any (except the first) larval instar before the last, and late diapause in the last instar. Early diapause was induced under a short day of 13 h or less at 28°C, and under both short and long daylength (12–16 h) at 20°C. The shorter the daylength and the lower the temperature, the younger the instar was entering early diapause. Early diapause was terminated by a long day (16 h) or a high temperature (28°C), after which larvae grew faster in short days than in long days until the last instar, when they again entered diapause, always in short days and frequently in long days. This late diapause was terminated also by an increase in daylength and was always followed by adult emergence. In this case, 13 and 14 h daylengths after exposure to 12 h daylength were as effective as 16 h daylength. Ourdoor samples collected in late autumn, winter and early spring at Hirosaki (40.5°N) comprised two distinct size groups, corresponding with the early and late diapause instars. Based on these results, the seasonal development strategy and intriguing aspects of the photoperiodic response in this cockroach are discussed.     

Dormancy, dispersal and the survival of cyclopoid copepods (Cyclopoida, Copepoda) in a lowland floodplain

1. The survival of cyclopoid copepods was investigated in a floodplain for 2 years where flooding occurred during the cold season. The cyclopoid community was studied in three waterbodies with distinct hydroperiods: a permanent pond connected to the flooded area during inundation, a temporary pool that is part of the flooded area and an isolated temporary pool.
2. Field studies, including data obtained from samples of water, sediment and soil, showed the overall predominance of species with a summer diapause over those with a winter diapause or without diapause. Emergence of cyclopoid copepods at the onset of flooding, examined using emergence traps and data from recently filled or still isolated temporary pools, showed that some species can survive several months of drying.
3. The ability of the diapausing fourth copepodid stages of Cyclops strenuus and C. insignis , the two cyclopoids most abundant during winter and spring flooding, to survive terrestrial conditions was tested in laboratory experiments. Both species survived for several months, but rates differed among the species. A higher percentage of C. strenuus survived for a longer period, possibly explaining why this species was relatively more abundant in more temporary habitats.
4. Both dormancy and dispersal facilitated survival of cyclopoid copepods in transient habitats connected to each other during flood periods. Dormancy was the most important survival strategy, whereas dispersal could be more important following prolonged periods without flooding.     

Biology and bionomics in Scotland of Anthocoris gallarum-ulmi

Abstract. 1. Observations were made on the biology of Anthocoris gallarumulmi De Geer (Hemiptera - Heteroptera: Anthocoridae) in West Central Scotland from 1973 until 1975. The life cycle was intimately linked with those of its principal prey species Schizoneura ulmi and S.patchae , leaf roll-gall aphids, and elm, their primary host ( Ulmus sp.).
2. Overwintered adults emerged in late April/May and could be found on a number of early flowering tree species, before congregating on elm in late May/June. This population was univoltine and exhibited obligatory female reproductive diapause.
3. Overwintered females emerged already mated but the subsequent pre-oviposition period was 25 days and oviposition period 27 days. Ova were deposited only in close association with galls of S.ulmi and Spatchae , and behavioural variations were shown between sites. Fecundity was c . 16 ova per female.
4. The incubation period was c . 8 days with the subsequent period of larval development 38–53 days, during which time the diet was almost exclusively either S.ulmi or S.patchae . Intergall migration was characteristic of post second instar larvae, which resulted in the concentration of fifth instar larvae and adults in a limited number of galls. It was during this period of local high population density that mating occurred.
5. Adults left elm within 14 days of imaginal ecdysis and thereafter, until overwintering, were recorded in only very low numbers from a range of tree, shrub and herb species.
6. Overwintering adults selected as hibernacula the bark of four tree species but principally Acer pseudoplatanus and Quercus robor . Females required a period of at least 75 days of cold 'shock' to terminate reproductive diapause.
7. Mortality among males surviving to spring emergence was 67%.     

Understanding gregariousness in a larval Lepidopteran: the roles of host plant, predation, and microclimate

Abstract.  1. Many moth and butterfly larvae are gregarious early in development, but become solitary in late instars. This ontogenetic variation in behaviour is probably the result of temporal changes in the costs and benefits associated with gregariousness. This study provides observational and experimental evidence that, in one particular moth species, a series of different ecological factors influence larval behaviour at different times during development.
2. Field observations show that young caterpillars of the limocodid Doratifera casta form large aggregations while foraging, but that mature larvae are largely solitary.
3. A field experiment revealed that individual first to third instar larvae in larger groups develop more rapidly, but that group size had no detectable influence on survival. The developmental advantage associated with gregariousness is affected by host plant species, but not by predator exclusion, suggesting that group living in these cryptic early instar larvae promotes feeding facilitation, but does not provide individuals with protection from natural enemies.
4. Laboratory experiments revealed that aposematic fourth instar caterpillars in large groups were less likely to be attacked by a generalist insect predator than those in small groups.
5. Field observations provided no evidence that group living affects body temperature, suggesting that microclimatic factors do not favour gregariousness in this species.
6. It is concluded that gregariousness in D. casta confers at least two different advantages on larvae at different stages early in development, but that these advantages disappear, or are outweighed by costs associated with intraspecific competition, in final instars.     

Oviposition date and pattern of embryogenesis in the grasshopper Chorthippus brunneus (Orthoptera, Acrididae)

The morphological development of eggs of the grasshopper Chorthippus brunneus , laid during successive periods through a single season, was closely monitored in the field at a site in NW England. Eggs within the same clutch developed synchronously. However, there were significant differences between clutches laid at the same time, reflecting spatial variation in microclimate (and possibly maternal effects and/or genetic differences). Eggs laid early in the season entered diapause and hatched relatively synchronously in the following spring. Eggs laid later failed to reach the diapause-stage before winter, but hatched, albeit late and with greater variance. Diapause was averted in these eggs. Plasticity in the nature of diapause has been documented from laboratory studies of many insect species (including C. brunneus ,) but this study directly relates the incidence of such developmental plasticity to oviposition date and seasonal temperature cycles in the field. The adaptive significance of diapause aversion and the evolutionary consequences for female reproductive behaviour are considered.     

Regulation of host diapause by an insect parasitoid

Abstract. 1. The interaction between larval development and parasitism by the braconid wasp Cotesia koebelei (Riley), was investigated in a population of the butterfly Euphydryas editha (Boisduval) (Nymphalidae). In this population, the butterfly host has an obligatory overwintering larval diapause.
2. It was found that E. editha larvae harbouring parasitoids were more likely to pass through an extra feeding instar before entering diapause than were non-parasitized conspecifics.
3. In addition, some individuals that were experimentally exposed to multiple parasitoid attacks bypassed diapause completely; these larvae passed through five or six feeding instars, reaching sizes typical of final instar post-diapause larvae.
4. The observed effect of superparasitism occurred regardless of whether the host larvae subsequently produced mature parasitoids, suggesting that parasitoid attack is sufficient to invoke the response.
5. It is proposed that the parasitoid C.koebelei regulates the number of pre-diapause feeding instars of its insect host E. editha, and that some component of the female venom, injected at oviposition, is responsible for this regulation.     

A trade-off between diapause duration and fitness in female parasitoids

Abstract 1. Diapause allows insects to cope with adverse weather conditions but also poses substantial fitness costs, e.g. through reduced survival of the diapausing stage or sublethal effects in the following season.
2. The energetic costs of diapause in females of the parasitoid Asobara tabida Nees (Hymenoptera: Braconidae) were assessed using experimental manipulation of diapause duration.
3. An increase in diapause length not only led to higher mortality among diapausing pupae but also caused a significant decrease in egg load, fat reserves, and dry weight of the emerging adult females.
4. Only larvae with sufficient resources were able to survive the entire diapause period and there was a trade-off between the metabolic costs of diapause and adult fitness components.
5. In contrast, the size (tibia length) of emerging females increased with increasing diapause duration. It is proposed that this may be due to a genetic correlation between larval dry weight and adult size, so that selection for heavier larvae would result in larger adults even when resources are limited.
6. Ultimately, the evolution of diapause duration and its consequences for adult performance will depend on the costs of overwintering and the ecological conditions in the habitat.     

Can the Antarctic terrestrial midge,Eretmoptera murphyi,tolerate life in water?

1. Early‐season flooding and ice entrapment at sub‐zero temperatures pose significant challenges to any polar terrestrial invertebrate. 2. The chironomid midge, Eretmoptera murphyi, is native to the sub‐Antarctic island of South Georgia and has been introduced to the maritime Antarctic (Signy Island). While the majority of its 2‐year life cycle is spent as a terrestrial larva, it is found in habitats potentially exposed to prolonged flooding. 3. The current study explored the tolerance of the larvae to extended submergence, demonstrating survival for at least 28 days, underlain by their ability to respire (oxy‐regulate) whilst submerged. To date, this ability is not known to be shared by any other terrestrial midge. Larvae also demonstrated notable anoxia tolerance whilst encased in ice, surviving for up to 28 days. 4. These data indicate a capacity to survive ecologically relevant periods of submergence and/or ice entrapment, such as may be experienced in their natural habitats.     

Factors affecting the rate of attack by Cotesia rubecula (Hymenoptera: Braconidae)

Abstract. 1. The relationship between responses of the insect parasitoid Cotesia rubecula (Marshall) to kairomones produced by the feeding activities of its host, Pieris rapae (L.), and patterns of parasitism were investigated under field conditions.
2. Parasitoid adults aggregated in patches with the highest densities of host larvae but there was no commensurate increase in the rate of attack in these patches.
3. The rate of attack was not limited by the availability of eggs.
4. The rate of parasitoid attack was highest where feeding damage by the host was highest, irrespective of current host density.
5. The rate of parasitoid attack was further influenced by host age distribution. Late instar larvae were less susceptible to parasitism than were early instar larvae. The rate of attack on early instar larvae occupying the same plants as late instar larvae was reduced. This reduction in rate of attack was due to limitations on parasitoid search time imposed by the increased feeding damage associated with large host larvae and by the increased time the parasitoid required to recover from an attack on these large host larvae.     

麦红吸浆虫滞育发生和解除过程中总脂和甘油三酯含量变化

【目的】脂类与昆虫滞育密切相关。本研究旨在明确麦红吸浆虫Sitodiplosis mosellana (Géhin)滞育过程中脂类物质含量的变化规律,探讨麦红吸浆虫滞育与脂类物质变化的关系。【方法】采用香兰素硫酸显色法测定了2008年5月－2010年1月不同时间陕西杨凌养虫圃内麦红吸浆虫滞育前、滞育期及滞育解除后幼虫和蛹体内总脂和甘油三酯含量。【结果】 滞育前幼虫总脂和甘油三酯含量分别为378.12和291.67 μg/mg,显著高于整个滞育期（P<0.05）。滞育年周期中,冬季总脂和甘油三酯含量最高,翌年1月结茧幼虫二者含量分别为335.29 和275.72 μg/mg,显著高于其他季节（P<0.05）;整个滞育期间,裸露幼虫和结茧幼虫总脂和甘油三酯随季节变化趋势相同,但同期结茧幼虫含量高于裸露幼虫;滞育当年与第2年同期幼虫总脂和甘油三酯含量差异不显著（P>0.05）。滞育解除后,总脂和甘油三酯含量随着幼虫的发育和变态逐渐降低,其中中蛹和后蛹显著低于活动幼虫（P<0.05）。【结论】麦红吸浆虫滞育不同时期幼虫及蛹总脂和甘油三酯含量存在明显差异,其滞育与脂类物质的含量密切相关。