Zur Artenfrage und zum Auftreten von Oscinella spp. in den Blütenständen wichtiger Getreidearten

Creation of the big trophical niches of the winter pine shoot moth (Rhyacionia buoliana Schiff.) on susceptible trees of Pinus silvestris L. provoked the mass outbreak of the pest in the south areas in Ukraine. In its turn, high density of the pest induced processes of the aggressivity's growth of population. In so doing the wide range of adaptive zones represented by groups of the pines with different resistance is favourable for formation of aggressive biotypes. The spreading of the pest from weakened to more resistant groups on the pines is similar to the process of “drift”.

This problem will be considered more detaily in the second part of this article.     

Genotype and environment interact to influence acceptability and suitability of white spruce for a specialist herbivore, Zeiraphera canadensis

Abstract.

• 1 The independent and interacting effects of plant genotype and site (i.e. environment) on the acceptability of white spruce, Picea gluaca (Moench) Voss, to the spruce bud moth, Zeiraphera canadensis Mut. & Free. and on plant suitability for egg development, were studied at four sites in New Brunswick, Canada.
• 2 A greater proportion of shoots on trees in two half-sib families, previously designated as highly susceptible, were partially eaten by spruce bud moths than shoots on trees in two half-sib families with low susceptibility.
• 3 At the site with the highest bud moth population, oviposition was highest on trees in susceptible families and on branches damaged by bud moth larvae. Oviposition was not higher on trees in susceptible families at the other three sites, resulting in a strong tree genotype × site interaction for oviposition.
• 4 Although there was a significant tree genotype × site interaction for egg predation, egg survival was higher on trees in susceptible families at all sites, due to lower levels of egg parasitism and predation.
• 5 Egg densities were positively but weakly correlated to shoot length and diameter. There were no consistent relationships between shoot length, shoot diameter, needle length or needle density and per cent egg survival, parasitism or predation.
• 6 Because egg survival was higher on trees in susceptible families at all sites but egg densities were only higher on trees in susceptible families at one site, host plant acceptability and suitability were positively related at only one site. We speculate that Z.canadensis does not distinguish between hosts of different suitability until trees are heavily damaged.

     

Effects of plant density on the survival rate of cabbage pests

The population density of herbivores depends on the spatial scale as well as the temporal scale. In a small-scale, short-term experiment, the number of individuals entering from the surrounding area will be most influential in determining the herbivore density. In large-scale, long-term experiments, however, the density of herbivores will rather be influenced by the survival rate of individuals inside the field because most of the herbivorous population derives from the parents that developed inside the field. If we want to predict the large-scale long-term density of herbivores, therefore, emphasis should be placed on the estimation of survival rate. To elucidate the effects of plant density on the large-scale long-term abundance of cabbage pests, we examined the survival rates of three lepidopterous pests, the small white butterfly Pieris rapae crucivora Boisduval (Pieridae), the beet semi-looper Autographa nigrisigna (Walker) (Noctuidae), and the diamondback moth Plutella xylostella (Linnaeus) (Yponomeutidae) under two levels of plant spacing (sparse plot, 2 m × 2 m interval; dense plot, 0.5 m × 0.5 m interval). The experiment with four blocks was repeated in two seasons. The number of eggs per plant was larger in the sparse plots than in the dense plots for all species. The survival rate of eggs and larvae, on the contrary, was lower in the sparse plots than in the dense plots. The lower survival rate of eggs in the sparse plots was mainly caused by the density dependency, while the lower survival rate of larvae in the sparse plots was mainly caused by the direct effects of plant density. It was thus suggested that the density of herbivores may become lower in the sparsely planted field in the long run because of the higher mortality of larvae. Received: September 16, 1998 / Accepted: March 22, 1999     

Intra- and interspecific density dependence mediates weather effects on the population dynamics of a plant–insect herbivore system

Temperature and precipitation are two major factors determining arthropod population densities, but the effects from these climate variables are seldom evaluated in the same study system and in combination with inter- and intraspecific density dependence. In this study, I used a 19 year time series on plant variables (shoot height and flowering incidence) and insect density in order to understand direct and indirect effects of climatic fluctuations on insect population densities. The study system includes two closely related leaf beetle species (Galerucella spp.) and a flower feeding weevil Nanophyes marmoratus attacking the plant purple loosestrife Lythrum salicaria. Results suggest that both intraspecific density dependence and weather variables affected Galerucella population densities, with interactive effects of rain and temperature on insect densities that depended on the timing relative to insect life cycles. In spring, high temperatures increased Galerucella densities only when combined with high rain, as low rain implies a high drought risk. Low temperatures are only beneficial if combined with little rain, as high rain cause chilly and wet conditions that are bad for insects. In summer, interactive effects of rain and temperature are different because high temperatures and little rain cause drought that induce wilting in plants, thus reducing food availability for the leaf feeding larvae. In contrast, the density of the flower feeding weevil was less affected by temperature and precipitation directly, and more indirectly interspecific density dependent effects through reduced resource availability caused by previous Galerucella damage.     

Non-specific resistance in brown planthopper resistant indica rice varieties against Plodia interpunctella (Lepidoptera: Phycitidae)

Mechanisms of host plant resistance against insect pests can be manifold. Resistance screenings generally use single target insect pests, but the resistance thus screened may not always be specific to the target insect species. We conducted a test for non‐specific resistance in indica rice varieties with resistance genes against brown planthopper (BPH), by using the Indian meal moth, Plodia interpunctella. The test system was very simple, and only required the non‐pest moth to be reared on rice flour. We compared the survival rate, developmental period and adult weight of the moth on three rice varieties: ‘Nipponbare’, a BPH‐susceptible japonica variety, and ‘Thai Collection 11’ and ‘Pokkali’, two resistant indica varieties. Our results were straightforward and demonstrate that resistance in the two resistant rice varieties is not BPH specific, because development of the moth was retarded and adult body weight was reduced.     

Population fluctuations of the diamondback moth,Plutella xylostella (L.) on cabbages inBacillus thuringiensis sprayed and non sprayed plots and factors affecting within-generation survival of immatures

Summary The population fluctuations and within-generation survival of immatures stages of the diamondback moth,Plutella xylostella on summer cabbages, were examined in unsprayed plots in 1984 and 1985 and in plots sprayed with two formulations ofBacillus thuringiensis in 1985. There was two distincy generations per crop and no noticeable difference in population fluctuations were observed in unsprayed plots between years. There was distinct difference between unsprayed and sprayed plots in 1985, from the larval period onwards. However, the survivorship patterns in all plots in both years were a Type 2 based onDeevey's (1947) classification. Life table studies showed that there was essentially no difference in the mortality agents acting on each of the stages except for numerical differences in the within-generation mortality rates. The major mortality rates during the egg stage were the parasitoid,Trichogramma spp. and unknown factors including rainfall; in the larval 1 stage was unknown mortality; in the larval 2 stage was the parasitoid,Apanteles plutellae and during the pupal stage was unknown mortality and parasitoid,Diadromus subtilicornis. Adult mortality was determined for generation 2. It was relatively higher in the unsprayed plots compared to theB. thuringiensis sprayed plots. The contribution of abiotic factors such as rainfall and temperature, and biotic factors such as parasitoids and predators in determining within-generation population levels and the fluctuation of populations on cabbage were discussed.     

Regulation of the seasonal population patterns of <Emphasis Type="Italic">Helicoverpa armigera</Emphasis> moths by <Emphasis Type="Italic">Bt</Emphasis> cotton planting

Transgenic cotton expressing the Bacillus thuringiensis (Bt) Cry1Ac toxin has been commercially cultivated in China since 1997, and by 2000 Bt cotton had almost completely replaced non-transgenic cotton cultivars. To evaluate the impact of Bt cotton planting on the seasonal population patterns of cotton bollworm, Helicoverpa armigera, the dynamics of H. armigera moths were monitored with light traps from four locations (Xiajin, Linqing and Dingtao of Shandong Province; Guantao of Hebei Province) in high Bt density region and five locations (Anci and Xinji of Hebei Province; Dancheng and Fengqiu of Henan Province; Gaomi of Shandong Province) in low Bt density region from 1996 to 2008. A negative correlation was found between moth densities of H. armigera and the planting years of Bt cotton in both high and low Bt density areas. These data indicate that the moth population density of H. armigera was reduced with the introduction of Bt cotton in northern China. Three generations of moths occurred between early June and late September in the cotton regions. Interestingly, second-generation moths decreased and seemed to vanish in recent years in high Bt density region, but this tendency was not found in low Bt density region. The data suggest that the planting of Bt cotton in high Bt density region was effective in controlling the population density of second-generation moths. Furthermore, the seasonal change of moth patterns associated with Bt cotton planting may regulate the regional occurrence and population development of this migratory insect.     

Plant regrowth response to a stem-boring insect: a swift moth-willow system

To examine plastic willow regrowth response to herbivory, we studied the effect of a boring insect, the swift moth Endoclita excrescens (Hepialidae: Lepidoptera), which does not remove apical meristems, on shoot growth in three willow species—Salix gilgiana, S. eriocarpa, and S. serissaefolia−by direct observations and experiments in the field. We hypothesized that the stem-boring could initiate new lateral bud activation, and result in secondary shoot regrowth without the removal of the primary apical meristems. There were significantly more lateral shoots on naturally attacked than unattacked stems, and a significant positive correlation between lateral shoot density and the number of swift moth tunnels per tree was observed for all three willow species. Artificial boring, and larval infestation, resulted in an increase in the number of lateral shoots, but did not affect growth of current-year shoots. The length of lateral shoots differed between species, being significantly longer in S. gilgiana than S. eriocarpa and S. serissaefolia. The results of this study show that compensatory regrowth can result even if herbivory does not remove the apical meristem. We argue that this type of plant compensatory response is probably widespread, given that the stem-boring is a common feeding type of insect herbivores.     

Intraspecific competition and subterranean herbivory: individual and interactive effects on bush lupine

High mortality of plants growing in dense monospecific stands (i.e. self-thinning) usually results from intense intraspecific competition. However, inconspicuous below-ground insect herbivory might be a potent but overlooked source of mortality within dense stands of plants, particularly if crowding limits a plant's ability to compensate for herbivore damage. Here I ask how high conspecific density influences a plant's ability to cope with heavy below-ground insect herbivory.
I manipulated conspecific density and exposure to an abundant root-borer, the ghost moth ( Hepialus californicus ), and examined the impacts on the fecundity, growth, and survival of bush lupine ( Lupinus arboreus ), a fast-growing shrub that grows in dense monospecific stands in coastal grasslands. Both herbivory and intraspecific competition affected seed production, size, and mortality of bush lupine over the two years of the experiment. Plants consistently produced fewer seeds when growing at high versus low density and ghost moth herbivory also significantly reduced seed production. The negative effects of herbivory on plant fecundity were similar, regardless of plant density. In contrast, plant survival was affected by both competition, herbivory, and the interaction of these factors. In high density plots, plant survival was uniformly low (averaging 0.45–0.50); plants exposed to herbivores died from heavy herbivory, and plants protected from herbivores died due to intense intraspecific competition that compensated for losses due to herbivory. In low density plots, ghost moth herbivory similarly reduced lupine survival, from an average survival probability of 0.94 in plots protected from these herbivores to 0.55 in plots exposed to herbivory. Thus, results show that regardless of plant density, below-ground herbivory can be a potent source of mortality.     

Specific impacts of two root herbivores and soil nutrients on plant performance and insect–insect interactions

Soil‐dwelling insects commonly co‐occur and feed simultaneously on belowground plant parts, yet patterns of damage and consequences for plant and insect performance remain poorly characterized. We tested how two species of root‐feeding insects affect the performance of a perennial plant and the mass and survival of both conspecific and heterospecific insects. Because root damage is expected to impair roots’ ability to take up nutrients, we also evaluated how soil fertility alters belowground plant–insect and insect–insect interactions. Specifically, we grew common milkweed Asclepias syriaca in low or high nutrient soil and added seven densities of milkweed beetles Tetraopes tetraophthalmus, wireworms (mainly Hypnoides abbreviatus), or both species. The location and severity of root damage was species‐specific: Tetraopes caused 59% more damage to main roots than wireworms, and wireworms caused almost seven times more damage to fine roots than Tetraopes. Tetraopes damage decreased shoot, main root and fine root biomass, however substantial damage by wireworms did not decrease any component of plant biomass. With the addition of soil nutrients, main root biomass increased three times more, and fine root biomass increased five times more when wireworms were present than when Tetraopes were present. We detected an interactive effect of insect identity and nutrient availability on insect mass. Under high nutrients, wireworm mass decreased 19% overall and was unaffected by the presence of Tetraopes. In contrast, Tetraopes mass increased 114% overall and was significantly higher when wireworms were also present. Survival of wireworms decreased in the presence of Tetraopes, and both species’ survival was negatively correlated with conspecific density. We conclude that insect identity, density and soil nutrients are important in mediating the patterns and consequences of root damage, and suggest that these factors may account for some of the contradictory plant responses to belowground herbivory reported in the literature.     

埃玛菌素对小菜蛾幼虫的毒力及其对子代种群增长的影响

研究了不同温度下埃玛菌素（emamectin）对小菜蛾Plutella xylostella幼虫的毒力,并用生命表法研究了25℃下埃玛菌素亚致死剂量LC₃₀）处理小菜蛾3龄初期幼虫后对其子代的影响。结果表明,埃玛菌素为典型的正温度系数药剂,对小菜蛾幼虫的触杀毒力在温度16～31℃间提高了10倍左右,口服毒力在相同温度范围内提高1 000倍左右,显示了埃玛菌素对该虫极高的口服毒力。25℃下埃玛菌素亚致死剂量处理当代幼虫后,对其子代的存活和繁殖均有明显影响,处理组和对照组子代的世代存活率分别为3.7%和35.5%,平均产卵量分别为92.4和148.3,种群趋势指数分别为1.8和27.4,净增殖率（R₀）分别为3.66和33.81,内禀增长力（r_m）分别为0.0665和0.0978。     

基于林分昆虫多样性调查的传统灯诱法改进效果分析

昆虫在生物多样性调查中具有不可替代的地位，灯诱法是效率最高的昆虫采集方法之一，然而传统的灯诱方法存在诱集范围无法界定、采样流程标准性不足等问题。为此，使用自制的聚光灯诱箱（Spotlight Box Trap，SBT）作为改进灯诱方法，并使用灯诱帐（Moth Collecting Tent，MCT）作为传统灯诱法进行对照；以北京杨（Populus×beijingensis）与油松（Pinus tabuliformis）人工林为样地，对两种灯诱装置采集的蛾类数量及物种组成、物种多样性指数与形态参数等多个指标进行评价。共采集蛾类24科164种6728号，其中北京杨林分SBT采集19科93种1164号，MCT采集22科125种2573号；油松林分SBT采集21科87种1044号，MCT采集22科117种1947号。分析显示：（1） MCT采集的个体数量与物种数量均高于SBT，总体上SBT在不同样地间的数据误差水平更低。（2）两种灯诱装置采集的蛾类优势种组成显著不同（P<0.05）；SBT共有种的数量比例始终高于MCT，即后者的特有种占比例更高。（3）总体上SBT物种多样性指数的数据误差水平更低。（4） MCT采集的蛾类物种体型更大（体长约为SBT的1.20倍）、飞行能力更强（相关参数约为SBT的1.20倍至1.36倍）。结果表明，在林分昆虫多样性的调查中，传统灯诱法的数据稳定性较差，对林分的代表性不足。聚光灯诱箱因其对光域的可控性和采集昆虫的规范性，具备更高的稳定性，在森林昆虫多样性调查与监测中具有更好的适用性与应用前景。     

Larger wildflower plantings increase natural enemy density,diversity, and biological control of sentinel prey,without increasing herbivore density

1. An important means of conserving beneficial insects in resource‐limited habitats is to meet their ecological requirements, which may be achieved by providing areas containing flowering plants that bloom throughout the season, but little is known about the importance of wildflower plot size for supporting natural enemies or the biological control they provide. 2. Wildflowers were established in plots of sizes ranging from 1 to 100 m², and found that natural enemy density, group richness, and diversity of natural enemy groups increased with plot size. 3. The density of insect herbivores was lower in all flower plots than in the control samples, whereas the diversity of herbivore groups was significantly higher in flower plots. 4. Comparing population growth of sentinel soybean aphids (Aphis glycines Matsumura) among plot sizes, aphid colonies were smaller as plot size increased. 5. Providing beneficial insects with flowering resources resulted in significantly more natural enemies and greater pest control than in smaller flower plots or mown grass areas. 6. These results indicate that the density, diversity, and function of natural enemies are sensitive to the size of wildflower plantings, even at relatively small scales. Therefore, larger wildflower plots are more suitable for the conservation of beneficial insects and their provision of natural pest control.     

Density-dependent resistance of the gypsy moth <Emphasis Type="Italic">Lymantria dispar</Emphasis> to its nucleopolyhedrovirus,and the consequences for population dynamics

The processes controlling disease resistance can strongly influence the population dynamics of insect outbreaks. Evidence that disease resistance is density-dependent is accumulating, but the exact form of this relationship is highly variable from species to species. It has been hypothesized that insects experiencing high population densities might allocate more energy to disease resistance than those at lower densities, because they are more likely to encounter density-dependent pathogens. In contrast, the increased stress of high-density conditions might leave insects more vulnerable to disease. Both scenarios have been reported for various outbreak Lepidoptera in the literature. We tested the relationship between larval density and disease resistance with the gypsy moth (Lymantria dispar) and one of its most important density-dependent mortality factors, the nucleopolyhedrovirus (NPV) LdMNPV, in a series of bioassays. Larvae were reared in groups at different densities, fed the virus individually, and then reared individually to evaluate response to infection. In this system, resistance to the virus decreased with increasing larval density. Similarly, time to death was faster at high densities than at lower densities. Implications of density–resistance relationships for insect–pathogen population dynamics were explored in a mathematical model. In general, an inverse relationship between rearing density and disease resistance has a stabilizing effect on population dynamics.     

Environmentally-based maternal effects: a hidden force in insect population dynamics?

Summary The nutritional environment of the parental generation of the polyphagous gypsy moth, Lymantria dispar, can significantly influence the growth and reproductive potential of the next generation through environmentally-based maternal effects. In the first experiment, members of the parental generation were reared on red oak trees (Quercus rubra L.) with known defoliation and phenolic levels. Diet in the offspring generation was homogeneous (synthetic diet). With genetic effects accounted for 1) offspring attained greater pupal weights when their mothers fed on trees with higher leaf damage levels, 2) daughters had a shorter prefeeding stage, a trait associated with dispersal tendency, when their mothers experienced higher condensed tannin levels, and 3) sons had lower pupal weights when their mothers experienced greater condensed tannin levels. In the second experiment, members of the parental generation were reared on either red or black oak (Q. velutina) trees. Offspring of each mother were divided among four diets: red oak, chestnut oak (Q. prinus L.), a standard synthetic diet, and a low-protein synthetic diet. The parental host species accounted for 24% of the variation in daughters' development time; offspring diet accounted for 52%. When mothers were reared on black oak rather than red oak, their offspring developed significantly faster when the F1 diet was chestnut oak. Environmentally-based maternal effects can significantly influence the expression of offspring dispersal potential, growth rate, and offspring fecundity. These traits contribute to natality and survival in natural populations and, hence, to population growth potential. Theoretical models of insect population dynamics demonstrate that the presence of a time delay in a density dependent response can induce destabilization. Maternal effects act on a time delay and may participate in the destabilization characteristic of outbreak species.     

Relative impact of spider predation and cover crop on population dynamics of Erythroneura variabilis in a raisin grape vineyard

Experimental and correlative evidence has steadily mounted over the past 30 years implicating spiders in the suppression of insect herbivore pests in crop fields. A large body of evidence has also shown that increasing agroecosystem vegetation diversity often influences the abundance of herbivores and their natural enemies. In previous experiments, the abundance of several species of spiders on grapevines in a raisin grape vineyard was twofold enhanced in vineyard plots vegetationally diversified with a cover crop. A concomitant reduction in the abundance of the leafhopper pest Erythroneura variabilis Beamer was observed on grapevines in the diversified plots, but a causal relationship was not established. In the present study, we simultaneously manipulated spider densities (in open‐vine spider exclusion and vine‐shoot enclosures) and ground cover to determine their relative impact on E. variabilis population dynamics. Open‐vine spider exclusion resulted in an average 35% increase in the density of E. variabilis the greatest effect with occurring during the first and second leafhopper generations. The negative impact of spiders on E. variabilis densities was corroborated with vine‐shoot enclosure experiments. Under the conditions of the present study, the cover crop per se did not affect the dynamics of E. variabilis populations on grapevines, despite a 1.6‐fold increase in spider densities on vines in cover crop plots, compared with vines in bare ground plots, probably due to insufficient spider enhancement and low overall E. variabilis abundance during the summer months. The cover crop had little effect on vine macronutrient status (and presumably vine water status). While this study provided further support for the hypothesis that vegetation diversity can enhance spider abundance, this enhancement does not always lead to lower pest densities, thus underscoring the complexity and variability that exists in interactions involving cover crop, spiders, and crop plants and their herbivore pests.     

Ramet size equalisation in a clonal plant,Phragmites australis

The influence of shading from older generations of dead culms (standing litter) on density, growth rate and development of size structure at the ramet level was investigated in a pure stand of Phragmites australis by experimental neutral shading of plots after removal of standing litter. Initial differences in height distribution between autumn and spring cohorts disappeared in the course of shoot growth. The Gini coefficients of shoot heights and estimated shoot weights indicated that the size structure of the shoots became more equal with increasing mean size in both shaded and unshaded plots. Relative growth rate for height (RHGR) and weight of individual shoots was negatively related to shoot size during the early and presumably storage-dependent growth period, suggesting a strong support for growth of smaller shoots. No etiolation was indicated by mean or maximum height in shaded and unshaded plots, or by the relationship between shoot height and weight. Mean shoot density was significantly lower in shaded than in unshaded plots in one of two shade treatment years. A regression model indicated a small but significant effect of shoot density on the approximately linear relationship between RHGR and the logarithm of height. The growth rate of small shoots was slightly larger at low than at high shoot density. Therefore, it is suggested that the shade from standing litter in P. australis stands can decrease shoot natality in the spring cohort, and thereby increase the support to fewer small shoots.     

Potential for controlling codling moth Cydia pomonella (Linnaeus) (Lepidoptera: Tortricidae) in Argentina using the sterile insect technique and egg parasitoids

Codling moth is the main pest affecting apples and pears worldwide. Most pest control strategies used against this insect have relied on the use of broad‐spectrum insecticides which have led to non‐desirable effects like pesticide resistance, residues in the environment, human health concerns and the reduction of access to international markets. Therefore, alternative pest control strategies that would result in sustainable fruit production systems while taking care of the environment are strongly promoted. The use of the sterile insect technique has proven to be a valuable pest control tactic within area‐wide integrated pest management strategies, and its synergistic effect for Lepidoptera pests when combined with other biological control tactics such as parasitoids has been documented. The purposes of this research were to evaluate the response of an Argentinean codling moth strain to a sub‐sterilizing radiation dose of 100 Gy and to assess the acceptability and suitability of sterile codling moth eggs by the egg parasitoids, Trichogramma cacoeciae (Marchal) and Trichogramma nerudai (Pintureau and Gerding). Irradiated female moths survived better than irradiated male moths and non‐irradiated male and female moths. Also, the fecundity of irradiated female moths was reduced by more than 30% as compared to non‐irradiated ones whereas their fertility was close to zero. The F₁ generation was male biased with a lower fertility (inherited sterility) than the parental generation. Trichogramma cacoeciae and T. nerudai parasitized both fertile and sterile eggs. However, there was a significant reduction in acceptability for sterile eggs. Trichogramma nerudai parasitized more eggs than T. cacoeciae, but egg acceptability for this species was proportionally lower than for T. cacoeciae especially on eggs oviposited by irradiated females. Development to adult of both parasitoids species was not substantially affected by the origin of the eggs and the wasps had acceptable levels of adult emergence, survival and fecundity. These results provided useful information on the potential for controlling the codling moth using egg parasitoids and the sterile insect technique in Argentina.     

Growth and survival of larvae of Thaumetopoea pinivora inside and outside a local outbreak area

1 Outbreaks of herbivorous insects tend to be spatially restricted, possibly because of demographic differences between inside and outside the outbreak area. In some cases, the margin of the outbreak area is distinct, allowing comparisons of adjacent areas that may identify factors leading to such differences in abundance. The northern pine processionary moth Thaumetopoea pinivora presently occurs at outbreak densities within a well‐defined area of approximately 3000 ha on the island of Gotland, south Sweden. We investigated whether cohorts of young larvae (first and second instar) had higher growth rate and survival inside or outside the outbreak area. 2 Group‐feeding appears to promote outbreaks in certain insect groups. Because T. pinivora larvae are highly social, we also compared larval performance between groups of different sizes inside and outside of the outbreak area: ‘small’ (33 eggs/group) and ‘normal’ (100 eggs/group). 3 Averaged over group size, whole colony mortality through the first two instars was two‐fold higher in the non‐outbreak area compared with the outbreak area. Mortality of individual larvae in the surviving colonies, however, did not differ between the two areas. There were only small differences in food quality (toughness, nitrogen content) between the areas, with no detectable effects on larval performance. 4 Larval relative growth rate did not differ between reduced and normal‐sized groups, which is surprising given that growth rate is known to increase with group size in other group‐feeding lepidopterans. 5 Reduced group size negatively affected larval survival, particularly in the outbreak area; by contrast, normal‐sized groups survived equally well in the two areas. Wood ants (Formica spp.) were more common outside the outbreak area, and appeared to be the main cause of colony mortality at low larval density. A different result was observed with regard to per‐capita mortality, which was higher in the outbreak area. We speculate that this could have been due to solitary predators being locally specialized on T. pinivora in the high‐density area.     

Statistical indicators for insects caught in the development trap

The effects of climate change on insect phenologies are of current interest, but little attention has been given to the potential driver for population decline of increased mortality arising from the failure to reach a viable overwintering stage. We use numerical simulation modelling to show that increased mortality from a partial second generation results in the weakening of density dependence, whereas increasing mortality within the first generation does not. These general results are then compared with annual trap count data for the moth Agrotis segetum (Denis and Schiffermüller) in Denmark, a species which is in decline there because a partial second generation fails to survive the winter. Results from the moth data are consistent with those from the simulations, and we conclude that the development trap phenomenon can be detected by the application of simple statistical tests to time‐series data.