Sexual size dimorphism in the two spot ladybird beetle Adalia bipunctata: developmental mechanism and its consequences for mating

Abstract  1. The literature on ladybirds indicates that males are consistently smaller than females but take the same length of time to complete their development. Rearing Adalia bipunctata at 20 and 25 °C confirmed that protandry cannot account for sexual size dimorphism in this species, nor can a difference in egg size.
2. Female larvae consumed more food and had a higher relative growth rate in the fourth instar than did male larvae.
3. When food is limited, small males appear to be more successful at mating than are large males.
4. To account for these results, it is hypothesised that the gonads of male larvae compete more strongly with the soma for resources and that this reduces the growth potential of the soma of male larvae relative to that of female larvae. The greater mating success of small males when food is limited supports the eat or mate hypothesis, which predicts that when food is limited small males will spend less time feeding and more time mating than will large males.     

Thermal ecology of western tent caterpillars Malacosoma californicum pluviale and infection by nucleopolyhedrovirus

Abstract 1. Western tent caterpillars hatch in the early spring when temperatures are cool and variable. They compensate for sub-optimal air temperatures by basking in the sun.
2. Tent caterpillars have cyclic population dynamics and infection by nucleopolyhedrovirus (NPV) often occurs in populations at high density.
3. To determine whether climatic variation might influence viral infection, the environmental determinants of larval body temperature and the effects of temperature on growth and development rates and larval susceptibility to NPV were examined.
4. In the field, larval body temperature was determined by ambient temperature, irradiance, and larval stage. The relationship between larval body temperature and ambient temperature was curvilinear, a property consistent with, but not necessarily limited to, behaviourally thermoregulating organisms.
5. Larvae were reared at seven temperatures between 18 and 36 °C. Larval growth and development increased linearly with temperature to 30 °C, increased at a lower rate to 33 °C, then decreased to 36 °C. Pupal weights were highest for larvae reared between 27 and 30 °C.
6. The pathogenicity (LD₅₀) of NPV was not influenced by temperature, but the time to death of infected larvae declined asymptotically as temperature increased.
7. Taking into account larval growth, the theoretical yield of the virus increased significantly between 18 and 21 °C then decreased slightly as temperatures increased to 36 °C.
8. Control and infected larvae showed no difference in temperature preference on a thermal gradient. The modes of temperature preference were similar to those for optimal growth and asymptotic body temperatures measured in the field on sunny days.
9. Warmer temperatures attained by basking may increase the number of infection cycles in sunny springs but do not protect larvae from viral infection.     

Superparasitism in Cotesia glomerata: response of hosts and consequences for parasitoids

Abstract.  1. Superparasitism occurs in Cotesia glomerata (Hymenoptera: Braconidae), a gregarious endoparasitoid of Pieris spp. (Lepidoptera: Pieridae). The response of P. brassicae larvae to superparasitism and the consequences for the parasitoid were examined in order to elucidate the ecological significance of this behaviour.
2. Field surveys of a Swiss population revealed that C. glomerata brood sizes from P. brassicae larvae ranged from three to 158, and both the female ratio and the body weight of emergent wasps correlated negatively with brood size. In the laboratory, single oviposition on P. brassicae larvae did not produce any brood size larger than 62, but brood size increased with superparasitism.
3. Laboratory experiments demonstrated that both naive and experienced female wasps were willing to attack hosts that had been newly parasitised by themselves or conspecifics. Superparasitism reduced survivorship but increased food consumption and weight growth in P. brassicae larvae. Superparasitism lengthened parasitoid development and prolonged the feeding period of host larvae.
4. Despite a trade-off between maximising brood size and optimising the fitness of individual offspring, two or three ovipositions on P. brassicae larvae resulted in a greater dry female mass than did a single oviposition on the host. Thus, superparasitism might be of adaptive significance under certain circumstances, especially when host density is low and unparasitised hosts are rare in a habitat.     

Influence of food quality and temperature on life history characteristics of the parthenogenetic mayfly, Cloeon triangulifer

SUMMARY. 1. Laboratory and field data indicate that Cloeon triangulifer McDunnough has at least three generations per year in White Clay Creek (Pennsylvania, U.S.A.).
2. The duration of the egg stage ranged from 5 days at 30°C to about 90 days at 10°C.
3. Larvae completed development (i.e. first instar to adult) in 27 days at 25°C, 45 days at 20°C, and 179 days at 10°C on an algal diet dominated by diatoms.
4. Larvae reared on hickory leaves completed development in 30 days at 25°C but died prior to metamorphosis at 10, 15 and 20°C.
5. Adult size (i.e. body length, wing length and dry mass) and fecundity were inversely related to rearing temperature for all laboratory and field experiments.
6. The significant interaction of food quality and temperature suggest that these factors may be important in understanding geographic variation in the life history of C. triangulifer.     

Natural thermal stress and heat-shock protein expression in Drosophila larvae and pupae

1. Whether Drosophila larvae and pupae naturally experience temperatures that can cause heat damage or death is poorly understood, but bears directly on numerous investigations of the thermal biology and heat-shock response in Drosophila . Accordingly, the temperatures of necrotic fruit, which Drosophila larvae and pupae inhabit, the temperatures of larvae and pupae outside the laboratory, and the levels of the heat-shock protein hsp 70 expressed by larvae in nature were examined.
2. When necrotic fruit was sunlit, internal temperatures rose to levels that can harm indwelling insects. Fruit size and evaporative water loss affected these temperatures. Temperatures of larvae and pupae in the field commonly exceeded 35 °C, with living larvae recorded at >44°C and pupae at >41°C. Natural mortality was evident, presumably because of heat.
3. In the laboratory, these temperatures kill larvae rapidly, with LT₅₀s (time taken for half the sample to be killed) of 30 min at 39 °C, 15 min at 40 °C and 8·5 min at 41 °C. Gradual transfer from 25°C to these temperatures resulted in no lesser mortality than did direct transfer.
4. Hsp 70 levels in lysates of whole larvae were measured by ELISA (enzyme-link immunosorbent assay) with an hsp 70-specific antibody. For larvae within necrotic apples experimentally transferred from shade to sun and within necrotic fruit in situ , hsp 70 levels equalled or exceeded levels detected in parallel laboratory studies of whole larvae or cells in culture.
5. These data provide an ecological context for studies of thermal stress and the heat-shock response in Drosophila that has heretofore been lacking.     

The relative importance of temperature and diet to larval development and adult size of the winter stonefly, Soyedina carolinensis (Piecoptera: Nemouridae)

SUMMARY. 1. Soyedina carolinensis Claassen, a leaf shredding stonefly, was reared in a series of three laboratory experiments from early instar to adult on different species of deciduous leaves and at various constant and fluctuating temperature regimes.
2. Experiment 1, which involved rearing larvae on fourteen different leaf diets at ambient stream temperatures, showed that diet significantly affected larval growth and adult size but did not affect overall developmental time.
3. Experiment 2, which involved rearing larvae on five different leaf diets at each of three fluctuating temperature regimes (viz ambient White Clay Creek (WCC), ambient WCC+3°C, and ambient WCC+6°C), showed that: (i) adding 6°C to the normal temperature regime of WCC was lethal to 99% of the larvae regardless of diet; and (ii) warming WCC by 3°C did not affect developmental time but did significantly reduce adult size relative to adults reared at WCC temperatures on certain diets.
4. Experiment 3, which involved rearing larvae on five different leaf diets at each of five constant temperatures (viz 5, 10, 15, 20, 25°C), showed that: (i) temperature significantly affected the mortality, growth, and development time of larvae whereas diet only affected larval growth and mortality; (ii) temperatures at or near 10°C yielded maximum larval growth and survival for most diets; (iii) at 5°C, larval mortality was high and growth was low resulting in a few small adults for most diets; (iv) larval mortality was at or near 100% at 15°C regardless of diet; and (v) no larvae survived at 20 and 25°C.     

Converse Bergmann cline in a Eucalyptus herbivore,Paropsis atomaria Olivier (Coleoptera: Chrysomelidae): phenotypic plasticity or local adaptation?

Aim  To measure latitude-related body size variation in field-collected Paropsis atomaria Olivier (Coleoptera: Chrysomelidae) individuals and to conduct common-garden experiments to determine whether such variation is due to phenotypic plasticity or local adaptation.
Location  Four collection sites from the east coast of Australia were selected for our present field collections: Canberra (latitude 35°19' S), Bangalow (latitude 28°43' S), Beerburrum (latitude 26°58' S) and Lowmead (latitude 24°29' S). Museum specimens collected over the past 100 years and covering the same geographical area as the present field collections came from one state, one national and one private collection.
Methods  Body size (pronotum width) was measured for 118 field-collected beetles and 302 specimens from collections. We then reared larvae from the latitudinal extremes (Canberra and Lowmead) to determine whether the size cline was the result of phenotypic plasticity or evolved differences (= local adaptation) between sites.
Results  Beetles decreased in size with increasing latitude, representing a converse Bergmann cline. A decrease in developmental temperature produced larger adults for both Lowmead (low latitude) and Canberra (high latitude) individuals, and those from Lowmead were larger than those from Canberra when reared under identical conditions.
Main conclusions  The converse Bergmann cline in P. atomaria is likely to be the result of local adaptation to season length.     

Effects of constant and variable temperature extremes on sex ratio and progeny production by Aphytis melinus and A.lingnanensis (Hymenoptera: Aphelinidae)

Abstract. 1. Aphytis melinus DeBach and A.lingnanensis Compere were exposed to temperature extremes during development, mating, preoviposition, and opposition to assess the effect of these exposures on the progeny per female and F₁ sex ratio.
2. Both species showed a significant reduction in the expected proportion of F₁ females when the male parent was exposed to 32°C during its development.
3. The proportion of females was further reduced when mating and/or oviposition occurred at 32°C and these effects on sex ratio appeared cumulative.
4. Preovipositional exposure of the females of both species to 38°C, 2°C or -2°C for 1 5, 4 or 7 h in most cases significantly reduced the expected proportion of females.
5. In general our results showed that the sex ratios of A.melinus was less severely affected by these exposures than were those of A.lingnanensis.
6. It is suggested that this difference may be one factor explaining the ability of A.melinus to exclude A.lingnanensis from the climatically more extreme areas of southern California.     

The responses of carrot fly larvae, Psila rosae, to components of their physical enrivonment

Abstract. 1. The responses of third instar Psila rosae (F.) larvae to light, temperature, humidity and soil moisture were investigated in the laboratory.
2. Larvae were photonegative and preferred a temperature of about 15°C. Temperatures between 30 and 40°C adversely affected movement and over 40° C were lethal.
3. In choice chambers, larvae preferred humidities of 70–100% r.h. and larvae in sand avoided dry conditions (2.5% field capacity). The latter response became more marked as larvae approached the pre-pupal stage when moistures of 40% field capacity and lower were avoided.
4. Most larvae were found at a depth of 8 cm in sand of uniform moisture content and temperature, but variation in moisture content could alter this preference.
5. In August, most larval damage in the field occurred near the tip of the carrot tap root but was more evenly distributed over the roots in November. It is uncertain whether this was due to soil near the surface being drier in August or whether it was caused by behavioural differences between the two generations of carrot fly larvae.
6. During the summer of 1975, low soil moisture levels resulted in the total absence of larval mines on the carrot roots even though pupae were found at depths of 20–30 cm in the soil. Temperature had no effect on the distribution of mines on carrot roots except at the top 2 cm of the soil profile.     

The roles of temperature and diapause in the life history of a temperate-zone dragonfly: Argia vivida (Odonata: Coenagrionidae)

Abstract. 1. The life cycle of Argia vivida Hagen generally took longer to complete in the field than was predicted on the basis of the thermal sum accumulated in laboratory rearing.
2. The prediction of a bivoltine life-cycle from geothermal sites with either a constant annual temperature of 26°C or thermal range of 11–31°C was not borne out because the intervention of short-day induced developmental delays in later larval instars extended the life cycle to 1 year.
3. This diapause, which synchronizes adult emergence with favourable summer temperatures, was also present in larvae from sites with annual temperature ranges of 0–33°C and 5–20°C.
4. At these colder sites completion of the life cycle takes 2 and 3 years respectively and dragonflies must be in cold-resistant stages during the winter. A long-day diapause, principally affecting late-instar larvae below a certain size during the summer, achieves this.
5. Large diurnal temperature fluctuations at the 0–33°C site markedly increase the useful thermal energy available to larvae for growth over that predicted by the thermal sum equation.
6. The interaction between the effects of temperatures favourable for growth and day-length-governed diapause, synchronize the emergence of the low-temperature sensitive adult stage of this tropical dragonfly with northern-latitude summers at a variety of habitats.     

An experimental study on the effects of salamander larvae on isopods in stream pools

SUMMARY 1. We used a combination of field surveys and field and laboratory experiments to examine the effects of predatory small-mouthed salamander larvae, Ambystoma barbouri (Kraus and Petranka), on the density and microhabitat use of benthic isopods, Lirceus fontinalis (Rafinesque).
2. A survey of four field stations yielded a significant negative relationship between the densities of salamander larvae and isopods.
3. A field experimental manipulation of salamander density confirmed that salamanders reduce isopod density and isopod use of open micro-habitat. Salamanders did not significantly affect the mean or coefficient of variation in isopod size.
4. A laboratory experiment showed that in the absence of refuges for isopods, salamanders were capable of eating large numbers of isopods; small isopods were particularly vulnerable to salamander predation.     

Influence of pregnancy on the thermal biology of the lizard, Sceloporus jarrovi: why do pregnant females exhibit low body temperatures?

1. Selected body temperatures of female lizards, Sceloporus jarrovi , were measured on a photothermal gradient during late pregnancy and again when postpartum, and pregnant females were subjected to one of three fluctuating temperature regimes that simulated body temperatures of (1) pregnant females, (2) postpartum females or (3) allowed normal thermoregulation.
2. Overall, females selected lower body temperatures when pregnant (mean = 32·0°C) than when postpartum (mean = 33·5°C).
3. Females regulated body temperature more precisely when pregnant than when postpartum as judged by their smaller variances in body temperature throughout the day.
4. When pregnant, females selected a lower mean maximum body temperature (mean: pregnant = 32·8°C; postpartum = 34·5°C) than when postpartum, but selected mean minimum body temperatures did not differ.
5. None of the experimental temperature treatments was detrimental to pregnant females. Female body length increased during pregnancy but the rate of increase did not differ among treatments. Moreover, length-adjusted body mass of postpartum females did not differ among treatments.
6. Pregnant females that experienced postpartum body temperatures produced neonates that were smaller in body mass and length than pregnant females that experienced pregnant body temperatures and females that were allowed to thermoregulate.
7. For neonates resulting from the postpartum body temperature treatment, the disparity in the body length, but not mass, was still observed at 9 days of age, although survival and growth of neonates was high and did not differ among treatments.
8. The results demonstrate that pregnant females could maintain higher postpartum body temperatures without compromising their physical condition, but select relatively low body temperatures, presumably to avoid decrements in offspring fitness.     

Depth selection by larval Ischnura verticalis (Odonata: Coenagrionidae): effects of temperature and food

SUMMARY. 1. Water temperatures in a shallow, weedy bay of a reservoir in southern Ontario were related to depth; mid-day temperatures in summer were approximately 30°C at the water surface and 20°C below 25 cm.
2. In the laboratory, larval Ischnura verticalis released in a thermal gradient (warmest water near the surface) did not remain near the surface any more than larvae in isothermal conditions. However, larvae in both isothermal and gradient conditions tended to spend more time close to the surface than expected by chance.
3. Larvae provided with food at all depths spent more time near the surface than did larvae with no food. Larvae provided with one feeding site in the coldest water remained near the food as much as did larvae provided with a single feeding site in the warmest water.
4. Results suggest that selection of microhabitats is based primarily on food availability and secondarily on proximity to the surface.     

Components of male fitness in relation to body size in Epirrita autumnata (Lepidoptera, Geometridae)

Abstract. 1. The effect of body size on different components of male fitness was studied for Epirrita autumnata , a geometrid known for its eruptive population dynamics. Body size is the main determinant of female fecundity in this species.
2. Longevity of males was found to have a weak negative correlation with body size at low temperatures. No significant correlation was found at higher temperatures.
3. We found no correlation between male size and female fecundity or egg size which is consistent with the small size of spermatophores in this species.
4. Small and large males were equally successful when allowed to compete for females in laboratory conditions.
5. In one or two field collections, males found mating were larger than males found singly. Large males also had an advantage in finding of virgin females, offered experimentally. No size-assortative mating was recorded.
6. We conclude that size-dependent mate location ability is the factor accounting for most of the variance in male fitness in E.autumnata. The dependence of fitness on body size may well be equally strong in males and females.     

Effects of elevated temperature on multi-species interactions: the case of Pedunculate Oak, Winter Moth and Tits

1. The effects of temperature on the Oak–Winter Moth–Tit food chain were studied at Wytham Wood, Oxford, and experimentally in the controlled environment solardomes at the Institute of Terrestrial Ecology, Bangor.
2. Tree cores from Wytham indicated that mature Oaks grew best at high temperatures and rainfall, but with low caterpillar populations. Young trees grew less well at elevated temperature, probably because they lost more water than they gained. Elevated temperatures advanced budburst, reduced foliar nitrogen and increased leaf toughness.
3. Moth eggs laid later or maintained at cooler temperatures than average required fewer heat units to hatch. Caterpillars took up to 50 days to complete growth at field temperatures but did so in only 20 days at a constant 15 °C.
4. The mass of Tit chicks at day 15 (day 1 = egg hatch) was positively correlated with temperature and negatively correlated with rainfall during the growing period.
5. At elevated temperature, budburst and moth egg hatch were synchronized, but earlier. Late feeding larvae and larvae fed on leaves from trees grown at elevated temperature produced smaller pupae. Pupal mass was unaffected when caterpillars and trees were maintained together under the same conditions.
6. Delaying egg hatch in Tits, to simulate conditions at elevated spring temperatures, resulted in reduced chick mass, body size and fledging success. This occurred because the chicks were fed later and prey quality was poorer, because the peak of caterpillar biomass was missed.
7. We predict that moth reproductive output will be retained at elevated temperatures because both leaves and caterpillars develop faster. Brood size in birds may be reduced because they cannot lay early enough to coincide with the narrower peak of food abundance.     

Size-dependent predation risk in tree-feeding insects with different colouration strategies: a field experiment

1. Body size is positively correlated with fecundity in various animals, but the factors that counterbalance the resulting selection pressure towards large size are difficult to establish. Positively size-dependent predation risk has been proposed as a selective factor potentially capable of balancing the fecundity advantage of large size.
2. To construct optimality models of insect body size, realistic estimates of size-dependent predation rates are necessary. Moreover, prey traits such as colouration should be considered, as they may substantially alter the relationship between body size and mortality risk.
3. To quantify mortality patterns, we conducted field experiments in which we exposed cryptic and conspicuous artificial larvae of different sizes to bird predators, and recorded the incidence of bird attacks.
4. The average daily mortality rate was estimated to vary between 4% and 10%. In both cryptic and conspicuous larvae, predation risk increased with prey size, but the increase tended to be steeper in the conspicuous group. No main effect of colour type was found. All the quantitative relationships were reasonably consistent across replicates.
5. Our results suggest that the size dependence of mortality risk in insect prey is primarily determined by the probability of being detected by a predator rather than by a size-dependent warning effect associated with conspicuous colouration. Our results therefore imply that warningly coloured insects do not necessarily benefit more than the cryptic species from large body size, as has been previously suggested.     

Reproductive success by large, closely related males facilitated by sperm storage in an aggregate breeding amphibian

The outcome of sexual selection on males may depend on female mate choice and male–male competition as well as postcopulatory processes such as cryptic female choice and sperm competition. We studied the outcome of sexual selection in the spotted salamander ( Ambystoma maculatum ), specifically examining the role of body size and relatedness on male reproductive success. Using controlled mating experiments in the field, we gave females access to three males of different sizes. We used seven microsatellite loci to determine paternity in the resulting larvae, estimate relatedness ( r ) between females and their mates, and calculate md ² (a measure of within-individual genomic divergence), heterozygosity, and standardized heterozygosity in the larvae. Both body size and relatedness to the female were significant predictors of male reproductive success. The relatedness of the males available to a female did not influence the amount of stored sperm she used to sire her larvae. Nonetheless, computer simulations showed that the average md ², heterozygosity, and standardized heterozygosity of the offspring were lower than expected by random mating. These differences are due to the use of stored sperm to fertilize some eggs; md ², heterozygosity, and standardized heterozygosity of larvae sired by stored sperm were significantly lower than those of larvae sired by the experimental males. These results suggest that relatedness may further influence a male's long-term reproductive success by determining whether his sperm is stored for later breeding seasons. Sexual selection in this salamander likely involves a complex interaction among many factors and may act over many seasons.     

Temperature-related fluctuations in the timing of emergence and pupation of Windermere alder-flies over 30 years

1. From 1966 to 1995, dates were recorded when adult alder-flies, Sialis lutaria L., were first seen (30-year range: 23 April – 25 May), 50% of the maximum density occurred (4 May – 4 June), and maximum density occurred (11 May – 17 June) along 200 m of Windermere shore. These emergence dates occurred at similar temperatures, estimated by mean values for both the emergence date and the week prior to emergence. The latter was the least variable at 10.1 °C (95% CL ± 0.37) for start of emergence, 11.2 °C (± 0.49) for 50% maximum density, 14.2 °C (± 0.51) for maximum density.
2. Final-instar larvae pupated in damp soil just above the water line. As laboratory temperatures were increased slowly from an initial 5 °C, the cumulative number of larvae leaving the water to pupate increased. A quadratic equation described this relationship from a threshold temperature of 7.2 °C to completion at 14.0 °C (50% point, 9.3 °C). The relationship between successful pupations and constant temperatures in the laboratory was well described by a quadratic equation with an optimum 14.9 °C (over 90% success) and no success outside the range 7–23 °C. A negative power-function described the relationship between days required for pupation and temperature, ranging from c . 28 days at 8.2 °C to c . 4 days at 22.1 °C.
3. Dates for larvae leaving the lake to pupate were back-calculated from dates for adult emergence, using the power-function for pupation time. Mean temperatures for estimated dates on which larvae left the lake to pupate were less variable than those for adult emergence, being 7.5 °C (± 0.20) for the start of pupation, 9.4 °C (± 0.16) for 50% maximum density, 13.7 °C (± 0.16) for maximum density. These values are similar to those obtained in the laboratory and can be used to predict pupation and adult emergence for different temperature regimes.     

Costs and consequences of superparasitism in the polyembryonic parasitoid Copidosoma koehleri (Hymenoptera: Encyrtidae)

Abstract.  1. Polyembryonic wasps provide dramatic examples of intra-specific developmental conflict. In these parasitoids, each egg proliferates into a clonal lineage of genetically identical larvae. If more than one egg is laid in a host (superparasitism), individuals of different clones may compete for food resources.
2. In the polyembryonic encyrtid Copidosoma koehleri , one larva per clone can differentiate into a sterile soldier. It is shown that soldiers are always females, and that they attack intra-specific competitors.
3. Research hypotheses were that (a) clones that develop in superparasitised hosts suffer heavier mortality than clones that develop in singly parasitised hosts, and (b) female clones cause higher mortality to their competitors than male clones, hence larval survival is lower in superparasitised hosts that contain females than in male-only broods.
4. The potential frequency of superparasitism in C. koehleri was manipulated by varying parasitoid–host ratios and exposure durations.
5. As parasitoid densities and exposure durations increased, the frequency of superparasitism rose, brood sizes increased, but the number of hosts that completed development was reduced. The number of offspring per parasitoid female decreased with increasing parasitoid–host ratios. Offspring size and longevity were inversely correlated with brood size. As superparasitism rates increased, fewer all-male broods were produced. Male–female broods were female-biased, suggesting selective killing of males by female soldiers. All-female broods were significantly smaller than all-male broods at high parasitoid densities only, possibly reflecting aggression among soldiers of competing clones.
6. The results support the working hypotheses, and suggest that female larvae outcompete males in superparasitised hosts.     

Survival at low temperature of larvae of the pine processionary moth Thaumetopoea pityocampa from an area of range expansion

1 Larvae of Thaumetopoea pityocampa (Lepidoptera: Notodontidae) develop throughout the winter, although their feeding activity and survival can be impaired by adverse climatic factors. The present study investigated the survival at low temperature of larvae originating from a population with range expansion in an alpine valley in Northern Italy.
2 The supercooling point of individually analysed larvae averaged at −7 °C. This value insufficiently described the cold hardiness of the larvae; 39% of the tested larvae were alive when returned to room temperature immediately after freezing. When larval colonies inside their nest were exposed to −17 °C for 1 h after gradual temperature decrease, survival was 70.4%.
3 Rearing of larvae in the laboratory at different day/night temperatures indicated an effect of cumulative chill injury on larvae. A logistic regression explained the relationship between negative thermal sum (h°C below 0 °C) received in the laboratory experiment and larval survival. A similar relationship was demonstrated between negative thermal sum and survival of larval colonies in the field.
4 In the laboratory experiment, some tested larvae were able to survive for up to 8 weeks without feeding depending on rearing temperature. As expected, feeding occurred only when larvae were reared at temperatures of 9 °C day/0 °C night.
5 We classify the larvae of T. pityocampa as being moderate freezing tolerant. The winter behaviour allows this species to track climate warming by a rapid expansion into those areas that become compatible with the insect's development.