Experiments on the responses of the larval stages of the geometrid moth Ectropis excursaria to light and gravity

The intensity-response curve to light in a vertical plane (where gravity was a conflicting stimulus to light) and a horizontal plane (where gravity was not a conflicting stimulus) suggests that the climbing behaviour of the first instar Ectropis excursaria (Guénée) is consistent with a summation of response to gravity and a response to light. Response to gravity was affected by the curvature of the crawling surface and the level of food deprivation of the caterpillars. On a 2 cm diameter crawling surface orientation to gravity occurred at an inclination between 20° and 30° from the horizontal. In a centrifugal field, a behaviour analogous to negative geotaxis occurred at accelerations of 3.5 ms^–2. Analysis of the geotactic turning tendencies of the caterpillar on flat inclined planes suggests that they exhibit meta-geotactic behaviour from inclinations of 40 to 90°. Negative geotaxis was exhibited by the caterpillars of later stadia although a response to gravity was lacking in the prepupal wandering caterpillars. The pre-pupal stage was also characterized by a reversal of the response to light.

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Résumé Un comportement d'escalade du premier stade d'E. excursaria correspondant à l'addition des réponses à la gravité et à la lumière, est suggéré par la courbe d'intensité de la réponse à la lumière, dans un plan vertical (où la gravité constitue un stimulus en opposition avec la lumière) et un plan horizontal (où la gravité n'est pas en opposition). La réponse à la gravité a été modifiée par la courbure de la surface à escalader et du degré de privation d'aliment des chenilles. Sur une surface d'escalade de 2 cm de diamètre, l'orientation par rapport à la gravité se produit entre 20 et 30° par rapport à l'horizontale. Das run champ de centrifugation, un comportement analogue à une géotaxis négative se produit pour une accélération de 3,5 m s^–2. L'analyse des tendances géotactiques des chenilles à tourner sur des plans inclinés suggère qu'elles présentent un comportement métagéotactique pour les inclinaisons de 40 à 90°. Des expériences d'immobilisation laissent penser que les antennes jouent un rôle dans la perception de la gravité. Les chenilles de stades ultérieurs ont présenté une géotaxie négative bien que la réponse à la gravité disparaisse chez les chenilles vagabondes en prénymphe. La prénymphose est aussi caractérisée par une inversion des réponses à la lumière.

     

Interactive effects of leaf maturation and phenolics on consumption and growth of a geometrid moth

Phenolic compounds are commonly regarded as the main chemical defenses of deciduous woody plants against insects. To examine how indices of leaf maturation (water content, toughness, and sugar/protein ratio) modified larval consumption and growth relative to phenolics and phenolic-related leaf traits, we measured consumption and growth of fourth-instar Epirrita autumnata (Bkh.) (Lepidoptera: Geometridae) larvae on three different days on young, normal, and mature leaves, respectively, from the same mountain birch (Betula pubescens ssp. czerepanovii (Orlova) Hämet-Ahti) trees. The larvae achieved the same growth rates on young and normal leaves, but had to consume 40% more on the latter. On more mature leaves, larval growth was poorer and was positively correlated with sugar/protein ratios (although the ratio peaked at that time). Indices of leaf maturation correlated with several phenolics in data pooled over the three study days, but poorly in any individual day. Similarly, in the pooled data, larval consumption and growth correlated with several leaf traits, but correlations between leaf and insect traits were few on any of the three days, and no trait was significant on each of the three days.We next examined whether variation in the maturation indices modified the associations of phenolics with insect consumption and growth. When interactions between phenolics and leaf maturation indices were taken into account, the number of phenolic compounds displaying significant associations with insect traits more than doubled. The relative importance of interactive versus direct associations increased with leaf maturation: on young leaves five phenolics showed direct and eleven interactive associations with insect traits, while in mature leaves we found two phenolics to display direct and thirteen phenolics interactive associations. Leaf water content, either alone or together with toughness and sugar/protein ratio, generally explained more of the variance in Epirrita growth (up to 59%) than any phenolic or phenolic-related trait alone (highest value 20%). Including interactive effects between phenolics and indices of leaf maturation in the model increased the proportion explained of variance in larval growth between 49 and 73%. Maturation indices explained 0 to 23% of variance in consumption, and the phenolic compound with the highest (positive!) correlation alone up to 28%, but taking into account interactions between phenolics and maturation indices raised the degree of explanation much (namely, 32 to 53%) over that explained by indices of leaf maturation alone. This indicates strong interactive effects on consumption between phenolics and indices of leaf maturation.     

The seed dimorphism of Spergularia marina in relation to dispersal by wind and water

Summary The seeds of the halophyte Spergularia marina differ both within and between individuals in that they either possess or lack a membranaceous border. This paper presents a morphological study of the length, weight and area of the seed types, and their dispersal characteristics under experimental conditions of wind and water dispersal. The winged seeds are shown to be larger both by length and by weight. Their rate of descent increases with wing loading. If the wing is lacking, however, the rate of descent increases with weight only. The distance of dispersal is equal for both seed types except at low wind speeds, when the winged seeds disperse farther. If the seed wing is removed, the excised seeds have shorter dispersal distances. When dispersed by water, a difference in the distance seeds are dispersed can only be detected in the presence of vegetation. The winged seeds are more frequently trapped in the vegetation as compared to the unwinged seeds. The hypothesis that the seed dimorphism is an adaptation for differential dispersal distances is discussed.     

Fluctuating asymmetry and survival ability in the forest tent caterpillar mothMalacosoma disstria: implications for pest management

Fluctuating asymmetry of the first tarsal segment of the proleg of the forest tent caterpiller mothMalacosoma disstria Hbn. (Lepidoptera: Lasiocampidae) was significantly inversely related to survival ability in the lab. The monitoring of population levels of fluctuating asymmetry could have important implications in pest management of this and other species by providing an indication of the health of a population.     

Observations of predator-prey interaction on dispersal of an oligochaete prey,Limnodrilus hoffmeisteri

Clumped patches of the oligochaete, Limnodrilus hoffmeisteri, were subjected to predation by the leech, Erpobdella punctata to determine how the dispersion pattern might change. Two separate experiments showed that worms left the patch in 39% and 48% greater numbers when the leech predator was present. This suggests that L. hoffmeisteri uses a antipredatory strategy of dispersal rather than clumping.     

荒漠孑遗植物裸果木种子时空扩散特性

裸果木(Gymnocarpos przewalskii)是亚洲中部荒漠区少有的第三纪孑遗物种,由于气候变化及人为干扰,其自然种群分布范围不断缩小。种子扩散作为植物生活史过程中的重要阶段,不仅对物种生存及其多样性至关重要,还影响物种分布范围和局部丰度。2015年和2016年分别在新疆哈密地区,采用布设种子收集器的方法,对其自然种群种子扩散的时空动态进行了定点连续观测。结果表明:该物种于当年6月上旬开始扩散,2015年略早于2016年。每年种子扩散持续时间约两个月,扩散趋势为单峰曲线,且呈集中大量扩散的模式,扩散高峰期与当年初次月降水高峰期吻合;在顺风的正南和东南方向上,种子扩散密度大且距离远;种子扩散主要集中在母株冠幅下,随着距母株距离的增加,种子扩散密度减少,二者间存在极显著的负相关性(P0.01),由于裸果木枝条繁多,对风力强度起到了一定的阻碍作用,可能是造成种子集中扩散在母株下的原因。裸果木种子扩散受外界环境(降水、风向)和自身因素等方面的影响,当种子在大量降水前完成扩散,将有利于种子在适宜的微生境萌发,是对多风、干旱的恶劣生境的一种长期适应。     

Life history and behavioural traits of Mnesampela privata that exacerbate population responses to eucalypt plantations: Comparisons with Australian and outbreak species of forest geometrid from the Northern Hemisphere

The larvae of the native Australian moth Mnesampela privata (Geometridae) sometimes defoliate plantation eucalypts, causing concern to industry. Typically, populations of M. privata maintain innocuous numbers in native forests, but outbreak populations can occur in plantations. Certain of the life history and behavioural traits of M. privata exacerbate population responses in simplified systems. Specifically, M. privata exhibits indiscriminate oviposition behaviour, for example, females oviposit upon leaves where conspecific egg clutches are already present. Combined with large egg clutches, this trait can lead to heavy exploitation of natal hosts. Complete defoliation of the natal tree necessitates that larvae disperse to undamaged hosts. In native forests, dispersing larvae would have a lower probability of locating a new host of suitable species and phenotype than they would in a plantation. Larval tolerance of a wide variety of species of Eucalyptus facilitates utilization of non‐natal eucalypts. A lower rate of encounter between larvae and natural enemies is achieved by means of leaf shelters and autumnal seasonal activity. Good dispersion by gravid females from plantations with large moth populations ensures that nearby plantations will be located. Given these characteristics, continued outbreaks of M. privata are likely in monospecific eucalypt plantations, especially when management practices reduce populations of natural enemies and overall vegetational complexity. However, because outbreaks only arise in plantations, population fluctuations of M. privata cannot be considered ‘self‐driven’. Hence, this moth is not a true eruptive species in the manner of other geometrids such as the autumnal moth (Epirrita autumnata autumnata) from Europe or the hemlock looper (Lambdina fiscellaria fiscellaria) from North America, but rather a gradient outbreak insect.     

Colonization of vineyards by phytoseiid mites: their dispersal patterns in the plot and their fate

The effect of wind and woody margins on the dispersal and population dynamics of phytoseiid mites was studied in a vine plot for a period of two years. Mites were sampled in the plot and in the surrounding vegetation (crops and natural vegetation) in order to determine phytoseiid mite abundance. The surrounding vegetation was considered to be a reservoir of phytoseiids from where the vine plot could be invaded. Directional and non-directional soil and aerial traps were placed in the plot to determine predatory mite exchange between the two areas. Colonization of the plot occurred in two stages: first, mite migration into the plot, followed by their establishment. The two-year study partially clarified the first of these two stages. Kampimodromus aberrans was the main species caught in the aerial traps. Phytoseiid mite dispersal within the vine plot seemed to be affected by both wind (direction, intensity and regularity) and phytoseiid mite density in the woody margin. However, the woody margin had a large effect only over a short distance. Some observations pointed towards an effect of other reservoir areas but it was not possible to characterize these. The population density of the phytoseiid mites in the plot increased from 1996 to 1998, but these increases are much smaller than one would expect on the basis of the number of mites migrating by air in the plot. Moreover, blocks where most mites were trapped were not the blocks where densities of phytoseiid mites on vine leaves were the largest. It therefore seems likely that not all migrants were able to develop. Their settlement pattern was not determined and this could constitute a potential research focus for the future.     

水流对莲草直胸跳甲种群存活和扩散的影响

【背景】水流是外来入侵生物传播蔓延的重要方式之一,特别是水生生物。水生杂草在水流传播过程中也经常携带其天敌从一个地方传播到另一个地方,如莲草直胸跳甲经常随着空心莲子草在水流中传播。研究水流对莲草直胸跳甲存活的影响及其对种群迁移的间接协助作用,可为解析该天敌抵御洪涝灾害和借助水流传播的能力提供科学依据。【方法】本研究以莲草直胸跳甲为研究对象,在室内进行水流模拟,比较水流转速（1000、1250和1750r.min-1）和时间（30、60、90和120min）对不同虫态莲草直胸跳甲（雌雄成虫、幼虫和卵）存活率的影响,并推算莲草直胸跳甲成虫和幼虫在野外随水流传播的最大距离。【结果】水流速度对昆虫的存活和发育具有一定影响。在不同旋转时间和转速下,雌雄成虫的存活率较高,幼虫次之,卵的存活率最低;雌虫随水流传播能达到的最远距离最大,雄虫次之,传播距离最小的是幼虫。【结论与意义】莲草直胸跳甲可随水流进行远距离传播和自然扩散,对于利用莲草直胸跳甲防治空心莲子草,缓解水渠和河道的堵塞非常有利。     

The complete nucleotide sequence of the mitochondrial genome of <Emphasis Type="Italic">Phthonandria atrilineata</Emphasis> (Lepidoptera: Geometridae)

Using long-polymerase chain reaction (Long-PCR) method, we determined the complete nucleotide sequence of the mitochondrial genome (mitogenome) of Phthonandria atrilineata. The complete mtDNA from P. atrilineata was 15,499 base pairs in length and contained 13 protein-coding genes (PCGs), 2 rRNA genes, 22 tRNA genes, and a control region. The P. atrilineata genes were in the same order and orientation as the completely sequenced mitogenomes of other lepidopteran species. The nucleotide composition of P. atrilineata mitogenome was biased toward A + T nucleotides (81.02%), and the 13 PCGs show different A + T contents that range from 73.25% (cox1) to 92.12% (atp8). Phthonandria had the canonical set of 22 tRNA genes, that fold in the typical cloverleaf structure described for metazoan mt tRNAs, with the unique exception of trnS(AGN). The phylogenetic relationships were reconstructed with the concatenated sequences of the 13 PCGs of the mitochondrial genome, which confirmed that P. atrilineata is most closely related to the superfamily Bombycoidea.     

中国尺蛾科新记录属——泽尺蛾属及其一新记录种记述（鳞翅目：尺蛾科：灰尺蛾亚科）

系统修订了中国绮夜蛾属的种类,将中国记录的11种订正为10种。其中,原记录的白缘绮夜蛾A.nivipicta经鉴定为大理石绮夜蛾A.marmoralis的误订名,迭绮夜蛾A.disrupta被鉴定为斑绮夜蛾A.sexpunctata的同物异名。检视标本均存放于中国科学院动物研究所标本馆。     

Analysis of odorant-binding proteins in antennae of a geometrid species, Ascotis selenaria cretacea, which produces lepidopteran Type II sex pheromone components

Information on the olfactory system in antennae of Geometridae moths is very limited, and odorant-binding proteins (OBPs) working as transporters of lipophilic odors have not been identified. In the first investigation on this family of insects, we examined antennal OBPs of the Japanese giant looper, Ascotis selenaria cretacea. RT-PCR experiments using several pairs of degenerate primers designed from known cDNA sequences encoding lepidopteran OBPs successfully amplified partial sequences of two pheromone-binding proteins (PBPs), named AscrPBP1 and AscrPBP2 in reference to their corresponding nucleotide sequence homologies with other PBPs. Using 5′- and 3′-rapid amplification of cDNA end strategies, a cDNA clone for AscrPBP1 encoding a protein of 141 amino acids was isolated. Western blotting with the antiserum against recombinant AscrPBP1 overexpressed in Escherichia coli showed that the AscrPBP1 gene was more strongly expressed in male antennae than in female antennae. Furthermore, natural AscrPBP1was isolated by immunoprecipitation with the antiserum, and its binding ability was evaluated by using synthetic sex pheromonal compounds with a C₁₉ chain. The result indicated that AscrPBP1 bound not only the pheromone components, 3,6,9-nonadecatriene and its 3,4-epoxy derivative, but also unnatural 6,7- and 9,10-epoxy derivatives. While no general odorant-binding proteins (GOBPs) were amplified in the RT-PCR experiments, two antisera prepared from GOBP1 and GOBP2 of Bombyx mori suggested the occurrence of at least two GOBPs in the A. s. cretacea antennae.     

Effects of weight loading on flight performance and survival of palatable Neotropical Anartia fatima butterflies

Previous studies show that the position of centre of body mass ( cm_body ) and the ratio of flight muscle to total body mass (flight muscle ratio, FMR) are good predictors of flight speed and manoeuvrability in butterflies. However, cm_body , FMR, and related morphometric traits are strongly correlated phcnotypically, making it difficult to identify the causal determinants of flight performance. By experimentally gluing weights that amounted to 15% body weight to a palatable Neotropical butterfly species (Anartia fatima) , we tested the effects of altering FMR and repositioning cm_body on two measures of flight performance: flight speed and the ability to evade capture. We then tested their effects on survival in a natural setting. Flight performance studies detected no significant differences in airspeed or evasive flight ability among unweighted controls, weighted-loaded butterflies (WL), and those with cm_body positioned further posterior (CM). In two mark-release-recapture experiments, survival of treatment groups did not differ, but males survived longer than females. In one experiment, WL and CM butterflies were recaptured more frequently than controls, whereas the probability of recapture for females was higher than that for males in the second experiment. When significant, results for recapture were consistent with a causal relationship between FMR and flight speed. Presumably, a decrease in flight speed was due to a reduction in muscle mass-specific power output in the weighted butterflies. However, the results did not support a relationship between manoeuvrability and cm_body     

Relative importance of ballistic and ant dispersal in two diplochorous Viola species (Violaceae)

Viola is one of the diplochorous plant genera that disperse their seeds in two ways, ballistic and ant dispersal. We compared the seed dispersal of two major Viola species of northern Japan, V. selkirkii and V. verecunda. The mean weight of seed was less in V. verecunda (0.42 ± SD 0.03 mg) than in V. selkirkii (0.61 ± 0.12 mg). The elaiosome of V. selkirkii (0.02 ± 0.004 mg) was larger than in of V. verecunda (0.006 ± 0.0004 mg), whereas the lipid component of elaiosome was not remarkably different between the two species. In ballistic dispersal, the mean dispersal distance was 56.0 ± 17.5 cm in V. verecunda but only 38.3 ± 5.1 cm in V. selkirkii. In ant dispersal, the mean dispersal distance was 28.1 ± 24.9 cm in V. selkirkii and 36.1 ± 33.7 cm in V. verecunda; however, the seed removal frequency of V. selkirkii (15.5%) was much higher than that of V. verecunda (3.0%). These results suggest that V. selkirkii is more dependent on ant dispersal while V. verecunda is more dependent on ballistic dispersal. The effect of seed predation was very serious in both species. In the quadrat census, 99.0% of V. selkirkii seeds and 99.1% of V. verecunda seeds were damaged by ground beetles, spiders, ticks, and others which frequently devoured diaspores. An experiment with V. verecunda seeds demonstrated that the overdispersion of seeds on the forest floor enhanced the frequency of removal by ants and reduced seed damage by predators.     

Influence of light and natural microbiota of the Butrón river on E. coli survival

The survival of an E. coli strain in water samples from the Butrón river has been studied. The input of E. coli cells in the aquatic system breaks down the established balance among the components of the natural microbiota: E. coli becomes the object of the active protozoal predation whereas the autochtonous heterotrophic community become alternative preys. As a result of this new situation, the natural microbiota increases but returns to the initial values once the E. coli cells have been removed from the system. The effect of the temperature of incubation on the survival is exerted through the effect of this parameter on the predatory activity of the protozoa. Light has a lethal and direct action on the E. coli cells, the effect of this parameter is even superior to that of predation.     

Effects of temperature, multiple allelochemicals and larval age on the performance of a specialist caterpillar

To understand the mechanisms underlying plant-insect herbivore interactions, it is necessary to examine the simultaneous effects of temperature, food quality and larval age. We examined the simultaneous effects of three allelochemicals (tomatine, rutin and chlorogenic acid) on the performance of first and second instar Manduca sexta larvae under two representative thermal regimes 21 : 10°C and 26 : 15°C for spring and summer, respectively. Thermal regime and allelochemicals interacted to influence the time from egg hatch to ecdysis to the third instar. On average, it took about half as much time to reach the third instar at 26 : 15°C as it did at 21 : 10°C. Separately, tomatine and rutin had a negative effect on developmental time from egg hatch to the third instar, but their simutaneous effects were not additive. Chlorogenic acid significantly reduced the negative effect of tomatine. The magnitude of the allelochemical effect was larger at the cooler thermal regime compared to the warmer regime. For instance, chlorogenic acid by itself had no effect at the 26 : 15°C regime, but at the 21 : 10°C regime it significantly shortened total developmental time. The effect of chlorogenic acid on stadium duration was distinctly different for the two instars. Chlorogenic acid shortened stadium duration of first instar larvae. However, depending on thermal regime and the presence of tomatine, chlorogenic acid had a negative, positive or neutral effect on stadium duration of second instar larvae. Molting duration of second instar larvae was shortened by a half day at the warmer thermal regime but was not affected by the allelochemicals. Final larval weight was influenced by rutin and chlorogenic acid. Caterpillars fed diets containing 20 moles of rutin were on average 10% lighter than those fed plain diet, whereas those fed diets containing 20 moles of chlorogenic adic were on average 7% heavier. However, the effect of chlorogenic acid depended on thermal regime. Overall, our results indicated that: 1) temperature and food quality can interact to influence insect performance and 2) these effects are influenced by larval age.     

Individual-based modeling of the spread of pine wilt disease: vector beetle dispersal and the Allee effect

Pine wilt disease is caused by the pinewood nematode Bursaphelenchus xylophilus, which is vectored by the Japanese pine sawyer beetle Monochamus alternatus. Due to their mutualistic relationship, according to which the nematode weakens and makes trees available for beetle reproduction and the beetle in turn carries and transmits the nematode to healthy pine trees, this disease has resulted in severe damage to pine trees in Japan in recent decades. Previous studies have worked on modeling of population dynamics of the vector beetle and the pine tree to explore spatial expansion of the disease using an integro-difference equation with a dispersal kernel that describes beetle mobility over space. In this paper, I revisit these previous models but retaining individuality: by considering mechanistic interactions at the individual level it is shown that the Allee effect, an increasing per-capita growth rate as population abundance increases, can arise in the beetle dynamics because of the necessity for beetles to contact pine trees at least twice to reproduce successfully. The incubation period after which a tree contacted by a first beetle becomes ready for beetle oviposition by later beetles is crucial for the emergence of this Allee effect. It is also shown, however, that the strength of this Allee effect depends strongly on biological mechanistic properties, especially on beetle mobility. Realistic individual-based modeling highlights the importance of how spatial scales are dealt with in mathematical models. The link between mechanistic individual-based modeling and conventional analytical approaches is also discussed.     

The interactive effects of competition and predation risk on dispersal in an insect

Dispersal dynamics have significant consequences for ecological and evolutionary processes. Previous work has demonstrated that dispersal can be context-dependent. However, factors affecting dispersal are typically considered in isolation, despite the probability that individuals make dispersal decisions in response to multiple, possibly interacting factors. We examined whether two ecological factors, predation risk and intraspecific competition, have interactive effects on dispersal dynamics. We performed a factorial experiment in mesocosms using backswimmers (Notonecta undulata), flight-capable, semi-aquatic insects. Emigration rates increased with density, and increased with predation risk at intermediate densities; however, predation had minimal effects on emigration at high and low densities. Our results indicate that factorial experiments may be required to understand dispersal dynamics under realistic ecological conditions.     

Evidence of partial anadromy and resident-form dispersal bias on a fine scale in populations of Oncorhynchus mykiss

We examine sympatric anadromous (steelhead) and nonanadromous (resident) rainbow trout (Oncorhynchus mykiss) from neighboring locations to test three hypotheses: (1) the sympatric life history types are not genetically different; (2) fine-scale dispersal is the same for both sexes, and (3) fine-scale dispersal is the same for steelhead and resident individuals. Data from 13 microsatellite loci reveal no genetic difference between sympatric steelhead and resident O. mykiss but moderate population structure (F _ST=0.019–0.028) between adjacent samples, regardless of life history type. Our results provide further evidence of partial anadromy and suggest that geographic proximity and genetic history, more than migratory type, should be considered when identifying populations for use in restoration of local genetic diversity. We find evidence of resident-form dispersal bias on a fine spatial scale, however, we find no evidence that fine-scale dispersal varies by gender. Conservation strategies should aim to maintain resident and anadromous forms when they occur in sympatry, as they may be important in facilitating gene flow on small and large spatial scales, respectively.     

The influence of constant and fluctuating temperatures on development rate and survival of pupae of the Australian sheep blowfly Lucilia cuprina

Functions describing instantaneous development rates in constant and natural temperature regimes were obtained for pupae of the Australian sheep blowfly, Lucilia cuprina (Wiedemann). These were derived using a technique that directly calculates rate functions from development-time observations made under any temperature regime. The functions indicated similar instantaneous development rates for constant and natural temperatures up to 30°C. At 30° the constant-temperature function reached a plateau which was maintained to the constant-temperature thermal limit. The natural-temperature function, however, continued its ascending phase to 34°, and then fell sharply to zero at 42°.The median survival temperatures of pupae for single 7-h exposures, daily 7-h exposures, and continuous exposures to high temperatures were 44.7, 39.2, and 34.4°, respectively. Development was completed at constant 15° but not at 10°. Median survival times at constant 10, 5, 0 and-5° were 4.3, 4.2, 2.5 and 1.5 days. Mortality was slight for single or daily 7-h exposures to-5°, but was complete for all except brief single exposures to-10°.

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Influence des températures constantes et de la thermopériode sur le taux de développement et la survie des pupes de Lucilia cuprina

Résumé Des fonctions associant les taux de développement aux températures en conditions constantes et en thermopériodes naturelles ont été établies pour les pupes de L. cuprina. La fonction pour les conditions naturelles décrit les taux de développement immédiat-c'est-à-dire les taux de développement existant en réponse à la température pour chaque laps de temps (en réalité, pour un court intervalle), plutôt que le taux moyen pour une gamme de températures. La fonction pour les températures constantes décrit à la fois les taux de développement instantané et moyen (ceux-ci étant équivalents, en ne supposant pas de réponses spécifiques suivant l'âge en conditions constantes). Les fonctions ont été dérivées en utilisant une technique qui calcule directement les fonctions des taux à partir des observations sur les durées de développement faites dans chaque condition de température.Les fonctions ont révélé des taux de développement instantanés semblables pour les températures jusqu'à 30°C. A 30°C la fonction pour la température constante a atteint un plateau qui s'est maintenu jusqu'à la limite thermique. La fonction pour les thermopériodes naturelles a continué de s'élever jusqu'a 34°C, et alors a chuté brutalement jusqu'à zéro à 42°C. Dans la région d'où les lots de pupes ont été obtenus les températures subies par les pupes dans les zones non ombragées dépassent fréquemment 50°C pendant l'été. Les taux de développement des pupes aux températures au dessus de 30°C, où les fonctions avec températures constantes et thermopériodes deviennent différentes, sont cependant cohérent avec les expériences dans la nature avec L. cuprina. Les températures médianes de survie des pupes pour une exposition unique de 7 h, pour des expositions quotidiennes de 7 h et pour une exposition constante à haute température étaient respectivement de 44,7; 39, 2 et 34,4°C. Par comparaison avec des températures estivales du sol de 50°C et plus, il semble vraisemblable que la mortalité nymphale est un facteur important de déclin des populations de L. cuprina au milieu de l'été.