The influence of ecological and life history factors on ectothermic temperature–size responses: Analysis of three Lycaenidae butterflies (Lepidoptera)

Body size has been shown to decrease with increasing temperature in many species, prompting the suggestion that it is a universal ecological response. However, species with complex life cycles, such as holometabolous insects, may have correspondingly complicated temperature–size responses. Recent research suggests that life history and ecological traits may be important for determining the direction and strength of temperature–size responses. Yet, these factors are rarely included in analyses. Here, we aim to determine whether the size of the bivoltine butterfly, Polyommatus bellargus, and the univoltine butterflies, Plebejus argus and Polyommatus coridon, change in response to temperature and whether these responses differ between the sexes, and for P. bellargus, between generations. Forewing length was measured using digital specimens from the Natural History Museum, London (NHM), from one locality in the UK per species. The data were initially compared to annual and seasonal temperature values, without consideration of life history factors. Sex and generation of the individuals and mean monthly temperatures, which cover the growing period for each species, were then included in analyses. When compared to annual or seasonal temperatures only, size was not related to temperature for P. bellargus and P. argus, but there was a negative relationship between size and temperature for P. coridon. When sex, generation, and monthly temperatures were included, male adult size decreased as temperature increased in the early larval stages, and increased as temperature increased during the late larval stages. Results were similar but less consistent for females, while second generation P. bellargus showed no temperature–size response. In P. coridon, size decreased as temperature increased during the pupal stage. These results highlight the importance of including life history factors, sex, and monthly temperature data when studying temperature–size responses for species with complex life cycles.     

Partial bivoltinism in a gregarious endoparasitoid: larval diapause as influenced by season and sharing a host

The yearly timing of the life cycle of a parasitoid is a key element of its life‐history strategy. I examine here factors influencing the expression of partial bivoltinism in Tetrastichus julis Walker (Hymenoptera: Eulophidae), a specialist parasitoid introduced to North America to attack its univoltine host, the cereal leaf beetle, Oulema melanopus (L.) (Coleoptera: Chrysomelidae). The varying tendency was assessed of individuals of this gregarious larval parasitoid to either emerge as adults in the same summer they mature, or to enter diapause to emerge the following year. Parasitized hosts were obtained by rearing cereal leaf beetles collected as mature larvae from grain fields in northern Utah (western USA) throughout the growing seasons in 2013 and 2014. Cocoons spun by these beetles were held to determine patterns over the spring and summer in the tendency of the parasitoid to forgo larval diapause. A high percentage (nearly 90% in 2013) of parasitoid individuals were found to forgo diapause and emerge in the same summer from earliest maturing hosts. This percentage rapidly declined to 20% or less of individuals forgoing diapause and emerging from cocoons as the summer advanced. The percentage of parasitoid individuals forgoing diapause increased significantly at a given time of season (early or late) as the number of conspecifics with which an individual shared a host larva increased. These results may reflect a trade‐off for individual parasitoids in which greater success in finding – and ovipositing in – host larvae the following spring vs. in summer, is countered by reduced survivorship in diapausing over the winter vs. emerging in the same summer in which the parasitoid matures. Expression of partial bivoltinism of T. julis, as affected strongly by both season and within‐host density, results in high rates of parasitism of cereal leaf beetles both early and late in the season.     

An exploration of genome size diversity in dragonflies and damselflies (Insecta: Odonata)

Like most insect orders, the Odonata (dragonflies and damselflies) remain poorly studied from the perspective of genome size. They exhibit several characteristics that make them desirable targets for analysis in this area, for example a large range in body size, differences in developmental rate, and distinct modes of flight – all of which are related to genome size in at least some animal taxa. The present study provides new genome size estimates and morphometric data for 100 species of odonates, covering about 1/5 of described North American diversity. Significant relationships are reported between genome size and body size (positive in dragonflies, negative in damselflies), and there is also indication that developmental rate and flight are related to genome size in these insects. Genome size is also positively correlated with chromosome number across the order. These findings contribute to an improved understanding of genome size evolution in insects, and raise several interesting questions for future research.     

SEASONALITY MAINTAINS ALTERNATIVE LIFE‐HISTORY PHENOTYPES

Many organisms express discrete alternative phenotypes (polyphenisms) in relation to predictable environmental variation. However, the evolution of alternative life‐history phenotypes remains poorly understood. Here, we analyze the evolution of alternative life histories in seasonal environments by using temperate insects as a model system. Temperate insects express alternative developmental pathways of diapause and direct development, the induction of a certain pathway affecting fitness through its life‐history correlates. We develop a methodologically novel and holistic simulation model and optimize development time, growth rate, body size, reproductive effort, and adult life span simultaneously in both developmental pathways. The model predicts that direct development should be associated with shorter development time (duration of growth) and adult life span, higher growth rate and reproductive effort, smaller body size as well as lower fecundity compared to the diapause pathway, because the two generations divide the available time unequally. These predictions are consistent with many empirical data. Our analysis shows that seasonality alone can explain the evolution of alternative life histories.     

Towards an interactive,process‐based approach to understanding range shifts: developmental and environmental dependencies matter

Many species are undergoing distributional shifts in response to climate change. However, wide variability in range shifting rates has been observed across taxa, and even among closely‐related species. Attempts to link climate‐mediated range shifts to traits has often produced weak or conflicting results. Here we investigate interactive effects of developmental processes and environmental stress on the expression of traits relevant to range shifts. We use an individual‐based modelling approach to assess how different developmental strategies affect range shift rates under a range of environmental conditions. We find that under stressful conditions, such as at the margins of the species’ fundamental niche, investment in prolonged development leads to the greatest rates of range shifting, especially when longer time in development leads to improved fecundity and dispersal‐related traits. However, under benign conditions, and when traits are less developmentally plastic, shorter development times are preferred for rapid range shifts, because higher generational frequency increases the number of individual dispersal events occurring over time. Our results suggest that the ability of a species to range shift depends not only on their dispersal and colonisation characteristics but also how these characteristics interact with developmental strategies. Benefits of any trait always depended on the environmental and developmental sensitivity of life history trait combinations, and the environmental conditions under which the range shift takes place. Without considering environmental and developmental sources of variation in the expression of traits relevant to range shifts, there is little hope of developing a general understanding of intrinsic drivers of range shift potential.     

Eggsac development rates and phenology of agrobiont linyphiid spiders in relation to temperature

Spider densities are often low after winter in annual crops, and crop management decimates spider populations several times per year. Population recovery rates and phenology depend on reproductive and development rates, which in turn are driven largely by temperature. We aimed to quantify the relationships between eggsac development rates and temperature in order to understand the relative value of different linyphiid species for the biological control of agricultural pests. Female adults of nine linyphiid species were collected from winter wheat in the UK over 3 years; Bathyphantes gracilis (Blackwall), Erigone atra (Blackwall), Erigone dentipalpis (Wider), Erigone promiscua O.P.‐Cambridge), Tenuiphantes tenuis (Blackwall) [formerly Lepthyphantes tenuis (Blackwall)], Meioneta rurestris (C.L. Koch), Oedothorax apicatus (Blackwall), Oedothorax fuscus (Blackwall), and Oedothorax retusus (Westring). These are agrobiont species that are dominant in agroecosystems. We tested how well development in the field can be predicted on the basis of laboratory experiments. We also built a simple phenology simulation model to test whether spider phenology in the field can be predicted by a general knowledge of the relationship between temperature and development rate. The relationships between temperature and development rates of eggsacs were not linear as described by a day‐degree model, but exponential as described by a biophysical model. Duration of the eggsac development period in the field was predicted accurately from laboratory experiments. We only found minor differences between development thresholds of eggsacs at constant temperatures compared with fluctuating temperatures. The phenology model predicted the phenology of L. tenuis and E. atra well, but the number of generations predicted for O. fuscus was not realised in the field. This suggests that development of this species may be affected by factors other than temperature. The methods used here could also be applied to other natural enemies, to determine whether their thermal biology is conducive to a role as biocontrol agents in disturbed agricultural systems.     

Control of the flexible annual/biennial life cycle of the heather psyllid Strophingia ericae

Abstract. Strophingia ericae (Curtis) (Homoptera: Psylloidea) takes one or two years to complete its life cycle. In both cases eggs hatch over a prolonged period from midsummer, possibly extending into the following spring at high altitude, and instars overlap in time. Instar III is the predominant overwintering stage in the lowland, annual cycle, whereas in the upland, biennial cycle most first-year nymphs overwinter in instars I and II and most second-year nymphs in instar V. When moved to the laboratory, instars IV and V from a predominantly annual population showed accelerated development in response to elevated temperatures and to long days in both mid-winter and early spring. In the biennial life cycle, short autumn days retard instar V development but the response to photoperiod disappears by the end of winter. Exposure to LD 18 : 6 h retarded development of early instars in the annual population, resulting in an accumulation in instar III. The proportion of overwintering instars I and II rises with increase in altitude and moult to instar III becomes progressively delayed. Nymphs that reach instar III under long daylengths in the year following hatching are channelled towards the biennial cycle. Exposure of a predominantly biennial population to 15 °C and LD 18 : 6 h after midsummer, thus avoiding autumn conditions, promoted the rapid development of overwintered nymphs, switching the cycle from biennial to annual.     

Evolution of alternative insect life histories in stochastic seasonal environments

Deterministic seasonality can explain the evolution of alternative life history phenotypes (i.e., life history polyphenism) expressed in different generations emerging within the same year. However, the influence of stochastic variation on the expression of such life history polyphenisms in seasonal environments is insufficiently understood. Here, we use insects as a model and explore (1) the effects of stochastic variation in seasonality and (2) the life cycle on the degree of life history differentiation among the alternative developmental pathways of direct development and diapause (overwintering), and (3) the evolution of phenology. With numerical simulation, we determine the values of development (growth) time, growth rate, body size, reproductive effort, adult life span, and fecundity in both the overwintering and directly developing generations that maximize geometric mean fitness. The results suggest that natural selection favors the expression of alternative life histories in the alternative developmental pathways even when there is stochastic variation in seasonality, but that trait differentiation is affected by the developmental stage that overwinters. Increasing environmental unpredictability induced a switch to a bet‐hedging type of life history strategy, which is consistent with general life history theory. Bet‐hedging appeared in our study system as reduced expression of the direct development phenotype, with associated changes in life history phenotypes, because the fitness value of direct development is highly variable in uncertain environments. Our main result is that seasonality itself is a key factor promoting the evolution of seasonally polyphenic life histories but that environmental stochasticity may modulate the expression of life history phenotypes.     

美国白蛾在我国发生的世代变化分析

美国白蛾是原产于北美地区的世界性植物检疫性害虫,自1979年发现入侵我国以来,严重威胁我国的生态安全。近年来,随着全球气候变暖,美国白蛾疫情不断南扩,其化性在一些地区出现新的变化。本文根据1979—2021年该虫发生扩散情况和各地区监测普查结果,结合部分地区温度变化数据,全面分析了美国白蛾入侵我国以来的发生扩散过程,对比分析了各地区发生世代数及变化情况,阐明部分地区世代数增加的原因。针对美国白蛾世代数演变规律和反复暴发成灾的情况,提出了当前应关注的几个重点问题:世代数增加引发的扩张速率显著上升;扩张潜力与沿长江地区扩散趋势减缓关系;暴发成灾时有发生与本土化趋势不明显。鉴于当前外来入侵物种管理要求提高及多年来美国白蛾防控工作中暴露的一些问题,提出要在守护国家生物安全底线前提下制定防治策略、在统筹生物多样性保护下实现控制目标、在做好应急准备下摸清暴发成灾规律、在强化疫情检疫管理中提高疫情认定效率等对策。