Delayed density-dependent parasitism of eggs and pupae as a contributor to the cyclic population dynamics of the autumnal moth

Many populations of forest Lepidoptera exhibit 10-year cycles in densities, with impressive outbreaks across large regions. Delayed density-dependent interactions with natural enemies are recognized as key factors driving these cyclic population dynamics, but emphasis has typically been on the larval stages. Eggs, pupae and adults also suffer mortality from predators, parasitoids and pathogens, but little is known about possible density relationships between mortality factors and these non-feeding life stages. In a long-term field study, we experimentally deployed autumnal moth (Epirrita autumnata) eggs and pupae to their natural enemies yearly throughout the 10-year population cycle in northern Norway. The abundance of another geometrid, the winter moth (Operophtera brumata), increased in the study area, permitting comparisons between the two moth species in predation and parasitism. Survival of autumnal moth eggs and pupae was related to the moth abundance in an inverse and delayed manner. Egg and pupal parasitoids dominated as density-dependent mortality factors and predicted the subsequent growth rate of the host population size. In contrast, effects of egg and pupal predators were weakly density dependent, and generally predation remained low. Parasitism rates did not differ between the autumnal and winter moth pupae, whereas predators preferred winter moth pupae over those of the autumnal moth. We conclude that parasitism of the autumnal moth by egg and pupal parasitoids can be related to the changes of the moth density in a delayed density-dependent manner. Furthermore, egg and pupal parasitoids cannot be overlooked as causal factors for the population cycles of forest Lepidoptera in general.     

Expansion of the winter moth outbreak range: no restrictive effects of competition with the resident autumnal moth

1. Both direct and indirect competition can have profound effects on species abundance and expansion rates, especially for a species trying to strengthen a foothold in new areas, such as the winter moth (Operophtera brumata) currently in northernmost Finland. There, winter moths have overlapping outbreak ranges with autumnal moths (Epirrita autumnata), who also share the same host, the mountain birch (Betula pubescens ssp. czerepanovii). Competitive interactions are also possible, but so far unstudied, are explanations for the observed 1–3 years phase lag between the population cycles of the two moth species. 2. In two field experiments, we studied host plant‐mediated indirect inter‐specific competition and direct interference/exploitation competition between autumnal and winter moths. The experimental larvae were grown either with the competing species or with the same number of conspecifics until pupation. Inter‐specific competition was judged from differences in pupal mass (reflecting lifespan fecundity), larval development time and larval survival. 3. Larval performance measurements suggested that neither direct nor indirect inter‐specific competition with the autumnal moth reduce the growth rate of winter moth populations. Winter moths even had a higher probability of survival when reared together with autumnal moths. 4. Thus, we conclude that neither direct nor indirect inter‐specific competition is capable of suppressing the spread of the winter moth outbreak range and that both are also an unlikely cause for the phase lag between the phase‐locked population cycles of the two moth species.     

Melanism in a larval Lepidoptera: repeatability and heritability of a dynamic trait

Abstract.  1. Although it is well established that the deposition of melanin pigment in the cuticle of larval Lepidoptera is influenced by both environmental and genetic factors, few studies have examined intra-individual regional variation in the degree of melanism or the ontogenetic dynamics of this trait. Here, heritable and density-dependent effects on within-individual and stage-specific variation in melanism were examined in caterpillars of the Egyptian cotton leafworm, Spodoptera littoralis (Boisduval).
2. Using quantitative spectrometric methods, it is shown that cuticular melanism changes dramatically within larval stadia, showing the highest and lowest levels of melanism early (first day) and late (final day) in each larval stadium respectively. However, solitary-reared caterpillars were significantly paler than those reared gregariously at all stages of development and maintained greater levels of variation in melanism. This variation in melanism was repeatable and exhibited a significant heritable component (narrow sense heritability based on offspring–parent regressions: h ² = 0.18–0.30).
3. The degree of melanism was correlated negatively with larval body weight in solitary caterpillars, but not gregarious ones. Melanism also varied spatially, with the lateral longitudinal band being consistently darker than the dorsal or dorso-lateral bands. Crowd-rearing increased melanism in all regions of larval cuticle, but the extent of crowding-induced melanism was more pronounced in the dorsal and dorso-lateral bands than in the lateral one.
4. These results indicate that although cuticular melanism is a highly dynamic trait, ontogenetic changes in relative cuticular melanism are both predictable and repeatable within individuals and genotypes. This has implications for our understanding of the evolution of melanism and for applying artificial selection on the basis of colour.     

Performance of the potentially invasive Siberian moth Dendrolimus superans sibiricus on coniferous species in Europe

1 The native range of the Siberian moth extends from the Pacific Ocean (Russian Far East, Japan and Northern Korea) across Siberia, Northern China and Mongolia to the Ural Mountains. At the beginning of the 21st Century, this species was documented west of the Ural Mountains in the Republic of Mari El, indicating range extension toward the west.
2 The Siberian moth has recently been suggested for regulation as a quarantine pest for European and Mediterranean Plant Protection Organization member countries. However, no specific report on European host plants for this pest has been published so far.
3 In the present study, larval host plant choice and performance was tested for the first time on coniferous tree species that are widely distributed and of commercial value in Europe.
4 Based on dual-choice tests on neonates and mortality, developmental duration and relative growth rates of the first- to sixth-instar larvae, we found European larch Larix decidua to be the most suitable host for the moth larvae, whereas European black pine Pinus nigra and Scots pine Pinus sylvestris were the poorest hosts. The remaining conifer species tested, European silver fir Abies alba , Nordmann fir Abies nordmanniana , and Norway spruce Picea abies , were intermediate host plants. Douglas-fir Pseudotsuga menziesii , originating from North America, was chosen by the larvae to the same extend as European larch, and was also highly suitable for larval development.
5 If the moth is introduced to European countries, it will become damaging in stands of European larch and Douglas-fir, mixed stands of fir and spruce; however, it will be less damaging in forests dominated by two-needle pines.
6 We predict that Dendrolimus superans sibiricus will be able to survive and develop on the main European coniferous tree species, including non-native coniferous tree species, resulting in severe damage to large areas of forests.     

Bottom-up and top-down effects in a tritrophic system: the population dynamics of Plutella xylostella (L.)–Cotesia plutellae (Kurdjumov) on different host plants

Abstract.  1. The effects of host-plant resistance on the population dynamics of the Diamondback moth, Plutella xylostella L., and its solitary parasitoid, Cotesia plutellae (Kurdjumov), were studied in replicated time-series experiments.
2. Host-plant resistance did not affect the equilibrial abundance of the Diamondback moth, but it affected the dynamics of Diamondback moth populations.
3. The mean population size of Diamondback moth showed no significant difference between Brassica rapa (a susceptible host plant) and Brassica napus (a partially resistant host plant) either in the presence or absence of the parasitoid.
4. Time-series analysis suggests that the dynamics of Diamondback moth on B. rapa were underpinned by delayed density-dependent processes. In contrast, the dynamics of the moth on B. napus were influenced by a direct density-dependent process.
5. Although measures of short-term parasitism showed a significantly higher rate of parasitism by C. plutellae on Diamondback moth feeding on B. napus compared with B. rapa , this individual performance does not translate into differences in the population dynamics. Analysis shows no significant difference in the persistence time of the population-level interaction between the host and parasitoid on the two different host plants.     

Fecundity of the autumnal moth depends on pooled geometrid abundance without a time lag: implications for cyclic population dynamics

1. The abundance and fecundity-related body size variation of the cyclic autumnal moth Epirrita autumnata were monitored from the early increase phase and throughout the outbreak to the end of the density decline in northernmost Norway during 1999-2006. Another geometrid, the winter moth Operophtera brumata, did not increase in density until the autumnal moth had its post-peak in 2004, and was at its own peak concurrent with the steeply declining autumnal moth abundance in 2005-06. 2. The body size variables measured (forewing lengths of males and females and hind femur lengths of males) of the autumnal moth showed a similar density-dependent response, i.e. increasing density was associated with decreasing body size and fecundity. Nevertheless, regression analyses clearly ranked the pooled geometrid abundance without a time lag as the best predictor for the body size variation, ahead of the abundance of the autumnal moth or past abundance of all geometrids. 3. Nondelayed effects of lowered food quality and absolute shortage of foliage under congested conditions are the most plausible reasons for reduced body size. 4. Two most commonly proposed causal factors of the autumnal moth population cycle, i.e. delayed inducible resistance of the host plant (mountain birch Betula pubescens czerepanovii) and delayed density-dependent parasitism by specialized hymenopteran parasitoids, cannot easily explain the diverging population trends between the autumnal and winter moths. 5. We suggest that either the inducible resistance of the host tree or the host utilization of the most important parasitoids and/or pathogens have to be strictly species-specific between these closely related moth species to produce the population dynamics observed. That fecundity of the autumnal moth was best related to the pooled geometrid abundance weakens support for the former hypothesis, while our lack of host-specific information limits conclusions about the role of natural enemies.     

Grape variety affects female but also male reproductive success in wild European grapevine moths

Abstract.  1. For insect herbivores the quality of the larval host plant is a key determinant of their fitness. Only little attention, however, has been given to the effects of plants on mating success of males and its consequence for the reproductive output of their mates. In addition, almost all the studies that have investigated the influence of host plants on herbivore fitness components have been done in the laboratory, and less is known of these effects in natural conditions.
2. Using the phytophagous European grapevine moth ( Lobesia botrana Den. & Schiff., Lepidoptera: Tortricidae), we tested the influence of grape cultivars as larval food on the probability of acquiring a mate for both sexes, and on the reproductive output of females and males.
3. Results from this study stress the importance of larval host plants on the reproductive success of both sexes. Larval diet differentially affected mating success and reproductive output of male and female moths. Fecundity, egg size, and egg hatchability were significantly different when larvae were fed on particular grape cultivars.
4. A given cultivar that is of poor quality for females is generally also of poor quality for males. A cultivar, however, could be suitable for females but not for males and vice-versa. Apparently, the nutrients required for adult reproduction are not necessarily the same for males and females.
5. The important conclusion from this study is that evaluating the differential effect of host-plant species on traits associated with reproductive success of herbivores requires that the effects on both sexes be taken into account.     

Influence of egg size differences within egg clutches on larval parameters in nine libellulid species (Odonata)

Abstract.  1. In libellulids, egg size differs between species and populations. There are also size differences within egg clutches of individual females.
2. Past experiments suggest that there are two different types of egg clutches in libellulids. Egg size decreases significantly during oviposition in species that perform non-contact guarding during oviposition. In contrast, in species ovipositing in tandem, egg size is randomly distributed.
3. This study deals with the possible consequences of egg size variation within the different egg clutch types. The study examined whether there is a correlation between egg development time, offspring sex or larval size and egg size.
4. The current experiments were conducted in Namibia and Germany. Five non-contact guarding and four tandem guarding libellulid species were used.
5. In some species larger eggs needed more time to develop, in some species no correlation between egg size and egg development time could be found, whereas in other species larger eggs developed faster.
6. The sex ratio was biased towards females in Leucorrhinia dubia and in Sympetrum striolatum and egg size was not associated with gender.
7. In both egg clutch types larger eggs resulted in larger larvae. In this study, evidence was found that the effects of egg size diminished with progressing larval development under good conditions. However, it is possible that the effects may have a greater influence under harsh circumstances.     

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.     

Low larval densities in northern populations reinforce range expansion by a Mediterranean damselfly

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Ecological mechanisms of population change during outbreaks of the spruce budworm

Abstract.  1. Stage-specific survival and recruitment of spruce budworm were measured by frequent sampling of foliage in four outbreak populations over a 15-year period in Ontario and Quebec, Canada.
2. Patterns of change in population density during the outbreak collapse phase were closely linked to changes in survival of the late immature stages, and were determined largely by the impact of natural enemies.
3. Host-plant feedback also contributed significantly to survival patterns throughout the outbreak: annual defoliation influenced survival of fourth and fifth instars and fecundity while cumulative defoliation influenced survival of the very early larval stages (first and second) via impacts on stand condition.
4. Inclusion of this host-plant feedback reveals spruce budworm population dynamics as a function of density-related trophic interactions that vary in their order and strength of influence over time. This view re-introduces the importance of forest interactions as a component of dynamics of the spruce budworm.     

What changed the demography of an introduced population of an herbivorous lady beetle?

1. The population dynamics of an introduced population of Epilachna niponica (Lewis) (Coleoptera: Coccinellidae) was investigated for a 7-year period following its introduction to a site outside of its natural range. A population from Asiu Experimental Forest was introduced to Kyoto University Botanical Garden, 10 km south of its natural distribution.
2. Arthropod predation was much lower in the introduced than in the source population. As a result of the lower predation in the Botanical Garden, larvae reached densities five times higher than in the Asiu Forest and host plants were frequently defoliated. Food shortage caused larval deaths from starvation and increased dispersal.
3. The density of the introduced population was much more variable than that of the source population. The variation in population density in both the introduced and source populations is limited by density-dependent reduction in fecundity and female survival. However, variation in the introduced population's density was increased due to host plant defoliation that resulted in overcompensating density-dependent mortality. In years with high larval density plants were defoliated and this increased adult mortality during the prehibernation period. Besides, the density-dependent regulatory mechanisms that produce population stability were weaker in the introduced population than in the source population.     

Snowshoe hare demography during a cyclic population low

1. Snowshoe hare ( Lepus americanus Erxleben) populations were studied in south-west Yukon during the low phase of the 10-year population cycle. Food availability and predator abundance were manipulated in a factorial design to determine the importance of each factor in hare dynamics during this phase.
2. Food was abundant during the low phase, and snowshoe hares were not food limited.
3. Survival of hares was higher than at any other phase of the cycle, and predators were scarce, but >75% of hare deaths resulted from predation.
4. Food addition resulted in higher hare densities and better body condition than on control sites. There were no observable effects of food addition on population rate of increase, recruitment, survival or age structure.
5. Mammalian predator reduction resulted in higher hare densities, higher survival, better body condition and an older age structure. Relative to control populations, recruitment was lower and population rates of increase similar.
6. The joint manipulation of food addition + predator reduction had greater positive effects on hare density and body condition than either single factor manipulation. Survival was better than on control sites, and the age structure was older than on control sites. Population rates of increase were similar, but recruitment was higher on the control areas.
7. We conclude that snowshoe hare dynamics at the low of the cycle are dominated by the interaction of food and predation. Risk of predation also had indirect effects on snowshoe hare age structure and body condition.     

Density-dependent effects of larval dispersal mediated by host plant quality on populations of an invasive insect

The success of invasive species is often thought to be due to release from natural enemies. This hypothesis assumes that species are regulated by top-down forces in their native range and are likely to be regulated by bottom-up forces in the invasive range. Neither of these assumptions has been consistently supported with insects, a group which includes many destructive invasive species. Winter moth (Operophtera brumata) is an invasive defoliator in North America that appears to be regulated by larval mortality. To assess whether regulation was caused by top-down or bottom-up forces, we sought to identify the main causes of larval mortality. We used observational and manipulative field and laboratory studies to demonstrate that larval mortality due to predation, parasitism, and disease were minimal. We measured the response of larval dispersal in the field to multiple aspects of foliar quality, including total phenolics, pH 10 oxidized phenolics, trichome density, total nitrogen, total carbon, and carbon–nitrogen ratio. Tree-level declines in density were driven by density-dependent dispersal of early instars. Late instar larvae dispersed at increased rates from previously damaged as compared to undamaged foliage, and in 2015 field larval dispersal rates were related to proportion of oxidative phenolics. We conclude that larval dispersal is the dominant source of density-dependent larval mortality, may be mediated by induced changes in foliar quality, and likely regulates population densities in New England. These findings suggest that winter moth population densities in New England are regulated by bottom-up forces, aligning with the natural enemy release hypothesis.     

Life-history strategy in an oligophagous tephritid: the tomato fruit fly, Neoceratitis cyanescens

Abstract.  1. In phytophagous insects, life-history traits mainly depend on host plant range. Substantial longevity, high fecundity and larval competition are the major traits of polyphagous Tephritidae while species with a restricted host range generally exhibit a lower longevity and fecundity as well as mechanisms to avoid larval competition. Our aim in this study was to investigate the life history of an oligophagous species, the tomato fruit fly, Neoceratitis cyanescens (Bezzi).
2. We determined life tables under laboratory conditions in order to calculate the main demographic parameters of N. cyanescens and studied the influence of larval and adult diet on life-history traits.
3. The mean longevity of N. cyanescens females was 40 days. There was a strong synchronisation of female maturity. Oviposition showed an early peak at 9–13 days after a short pre-oviposition period (6 days). The absence of proteins in the adult diet both delayed ovarian maturation and decreased female fecundity. In addition, females originating from tomato fruits produced significantly more eggs than females originating from bugweed or black nightshade, showing that even the larval host plant may strongly affect the subsequent fecundity of adult females.
4. The traits of N. cyanescens are then discussed in the light of those documented for polyphagous and monophagous tephritids. Neoceratitis cyanescens displayed attributes intermediate between those of polyphagous and monophagous tephritids. Its smaller clutch size compared with polyphagous species and its specialisation on the Solanaceae family whose fruits contain toxic compounds may help in reducing intra- and inter-specific competition, respectively.     

The Lepidoptera fauna associated with Calluna vulgaris: effects of plant architecture on abundance and diversity

Abstract .1. The effects of plant architecture on the Lepidoptera assemblage associated with heather Calluna vulgaris were studied at four locations in northern England and southern Scotland in 1992 and 1993. The study areas were Calluna vulgaris – Vaccinium myrtillus heaths, where management by rotational burning had created a mosaic of stands of different-aged Calluna . Lepidoptera were sampled in the larval stage.
2. Larval abundance was considered in relation to Calluna age, height, cover, green shoot density, and flower density. For most study areas and species, multiple regression selected Calluna height as the variable that explained most variation in larval density, although the explanatory variables were intercorrelated.
3. After compensating for the effects of different study locations, the slope of the regression between the logarithm of larval density and the logarithm of Calluna height was common to the macrolepidoptera, microlepidoptera, geometrid, and noctuid larval groups.
4. There was a significant progressive increase in larval diversity with increase in Calluna height, due to the presence of uncommon moth species in the samples from taller Calluna, and a change in the contribution of common species to the community in different height zones.
5. The significance of these results is discussed in relation to other heathland invertebrate studies and management practices.     

Migration and Allee effects in the six-spot burnet moth Zygaena filipendulae

Abstract 1. Migration into local populations may increase the likelihood of persistence but emigration may decrease the persistence of small and isolated populations. The dispersal behaviour of a day-flying moth Zygaena filipendulae was examined to determine whether emigration is correlated positively or negatively with population size and host plant density.
2. A mark–release–recapture study showed that most moths moved small distances (< 40 m on average) and only 6% of movements were > 100 m.
3. Twenty-five individuals moved between populations, a measured exchange rate of 8%. Moths were more likely to move between patches that were close together and they moved to relatively large patches.
4. The fraction of residents increased with increasing population size in the patch and increasing host plant cover. Relatively high proportions of individuals left small patches with small moth populations.
5. Moths released in grassland lacking Lotus corniculatus (the host plant) tended to leave the area and biased their movement towards host plant areas, whereas those released within an area containing L. corniculatus tended to stay in that area.
6. Biased movement away from small populations and areas of low host plant density (normally with low population density) was found. This migration-mediated Allee effect is likely to decrease patch occupancy in metapopulations, the opposite of the rescue effect. The effects on metapopulation persistence are not known.     

Long-term responses of zooplankton to invasion by a planktivorous fish in a subarctic watercourse

1. Introduced or invading predators may have strong impacts on prey populations of the recipient community mediated by direct and indirect interactions. The long-term progression of predation effects, covering the invasion and establishment phase of alien predators, however, has rarely been documented.
2. This paper documents the impact of an invasive, specialized planktivorous fish on its prey in a subarctic watercourse. Potential predation effects on the crustacean plankton, at the community, population and individual levels, were explored in a long-term study following the invasion by vendace ( Coregonus albula ).
3. Over the 12-year period, the density and species richness of zooplankton decreased, smaller species became more abundant and Daphnia longispina , one of the largest cladocerans, was eliminated from the zooplankton community.
4. Within the dominant cladocerans, including Daphnia spp., Bosmina longispina and Bosmina longirostris , the body size of ovigerous females and the size at first reproduction decreased after the arrival of the new predator. The clutch sizes of Daphnia spp. and B. longirostris also increased.
5. Increased predation pressure following the vendace invasion induced many effects on the crustacean zooplankton, and we document comprehensive and strong direct and indirect long-term impacts of an introduced non-native predator on the native prey community.     

Microclimate and tree community linked to differences in lepidopteran larval communities between forest fragments and continuous forest

Aim  We aim to assess the impact of forest fragmentation on lepidopteran larval community and study the associations of microclimate and tree community with lepidopteran assemblage.
Location  Kibale National Park, Uganda.
Methods  We investigated the effects of forest fragmentation on leaf herbivory, density of lepidopteran caterpillars, species richness and diversity as well as the composition of herbivorous lepidopteran larval community. Microclimate, size of the fragment, distance to the continuous forest, and tree diversity were studied as possible explanatory factors. We sampled 10 Neoboutonia macrocalyx Pax. (Euphorbiaceae) trees in each fragment during dry and rainy season, total of four times, in a year to cover the seasonal variation.
Results  The rates of herbivory, total larval density and species richness were significantly lower in the forest fragments than in the continuous forest but species diversity expressed as Fisher's alpha did not differ. The dominance structure and community composition of the larval communities in the fragments was different from that of the continuous forest. None of the differences we observed were related to the fragment area or distance to the continuous forest. Instead, we found an indication of association between the herbivore and the tree communities. The fragments had significantly lower humidity during most of the day and higher temperature during the afternoons (14–17 h), which might partially explain the differences in lepidopteran larval communities.
Main conclusions  Decreased larval density and species richness as well as differences in the community composition and structure all highlight the importance of large continuous forest areas for maintaining larval biodiversity.     

Oviposition in an eruptive moth species, Yponomeuta evonymellus, is insensitive to the population density experienced during the larval period

We studied if the population density experienced during larval development affects the reproductive schedule of Yponomeuta evonymellus L. (Lepidoptera, Yponomeutidae), a moth species characterized by outbreak population dynamics. More specifically, we predicted that reproduction would be delayed to facilitate emigration from sites with suboptimally high densities of conspecifics. We manipulated larval densities in the laboratory, as well as in those pupae collected from the extremes of natural densities. As the response, we also recorded the timing and sizes of egg clutches laid. The results did not support our initial predictions: the timing of oviposition was not dependent on larval growth conditions. This apparent lack of adaptation might be related to the loss of ‘memory’ during metamorphosis in holometabolous insects. However, oviposition schedules were also only minimally sensitive to elevated adult density. An inability to respond to high larval densities may contribute to the outbreak dynamics in this species.