Moth outbreaks in relation to oak masting and population levels of small mammals: an alternative explanation to the mammal-predation hypothesis

The relationship between population outbreaks of the gypsy moth (Lymantria dispar) and oak masting in North America has been interpreted as an effect of reduced predation on moth pupae from small mammals after years of acorn failure. However, moth defoliation could be a consequence of high acorn production rather than of acorn failure, as all moth outbreaks in two time series presented by Liebhold et al. [Popul Ecol (2000) 42:257–266] were reported shortly after mast years. A similar pattern has been found for the green oak leaf roller moth (Tortrix viridana) in southern Norway. Because predation from small mammals should be less important for the latter species, I argue that the acorn-moth relationships are most likely caused by mast-induced changes in the chemical composition of oak leaves. Given the high number of eggs laid by each moth female, there is a huge potential for population growth in or shortly after a mast year if larval survival is no longer limited by low food quality.     

Dormant Season Prescribed Fire as a Management Tool for the Control of <Emphasis Type="Italic">Salix caroliniana</Emphasis> Michx. in a Floodplain Marsh

Expansion of woody species into herbaceous wetlands is a serious concern in wetland management. Prescribed fire is often used as a tool to manage woody species, although many species resprout after fire making control problematic. In this study, we assessed the usefulness of repeated dormant season fires for controlling Salix caroliniana (Michx.) in a floodplain marsh in Florida. Salix is a common shrub in southeastern marshes that resprouts prolifically after fire. We compared stem basal area, stem density, and cover of Salix in three adjacent sites in a floodplain marsh in east central Florida. One site was burned once in February 1997, another site was burned in February 1997 and then again in March 1999 and one site was left unburned. At the unburned site, Salix stem basal area, stem density, and cover increased over the course of the study. In the two burned sites, the first fire destroyed large diameter stems and stimulated production of sprouts. As a result, stem basal area and cover decreased but stem density remained unchanged. The second fire caused a decline in stem density and a further decline in cover. Changes in understory species composition and cover could not be attributed to the fires. Our results suggest that dormant season fires are effective in reducing Salix cover and basal area, and that repeated fires have greater effects than a single fire.     

Adaptation of the gypsy moth to an unsuitable host plant

The pattern of adaptation with regard to life history traits and traits thought to be important in feeding habits of caterpillars in two populations of the gypsy moth (Lymantria dispar L.; Lepidoptera: Lymantriidae) originating from the locust tree (Robinia pseudoacacia; Fabaceae) and oak (Quercus petrea; Fagaceae) forests were investigated in the laboratory. The Robinia population has experienced unsuitable locust tree leaves as an exclusive food resource for more than 40 years. Since Quercus species are the principal host plants of the gypsy moth, the specific objectives of this study have been to measure the extent of differentiation between ancestral and derived populations in several life history traits (egg-to-adult viability, duration of larval and pupal stages, and pupal weight) and nutritional indices – relative growth rate (RGR), relative consumption rate (RCR), assimilation efficiency (AD), gross growth efficiency (ECI), and net growth efficiency (ECD). Significant differences between the Quercus and Robinia populations were detected in pupal duration, RGR, RCR, and AD. The presence of a significant population × host interaction in traits such as preadult viability, duration of pupal stage, RGR, and ECI suggests that adaptation of the gypsy moth to the unsuitable host might be ongoing. Using a full-sib design, we screened for genetic variation in life history traits within both populations, and examined the genetic correlations of performance across oak and locust leaves within both populations. The genetic variances for analyzed life history traits were lower under conditions that are commonly encountered in nature. Our data show that positive cross-host genetic correlations preponderate within both populations.     

Why the larch bud-moth cycle collapsed in the subalpine larch-cembran pine forests in the year 1990 for the first time since 1850

The larch bud-moth cycle has been observed in the sub-alpine larch-cembran pine forests 16 times since 1850. Infestation is easily recognized by the characteristic red-brown discoloration of the larch crowns due to the wasteful feeding of the bud moth larvae. The heaviest defoliation recurs at intervals of 8.47±0.27 (SE) years, and the larval density per kilogram of larch branches varies more than 10000-fold over four or five generations. The basic regulatory mechanism for this cycle is the induced change in food quality for the two or more subsequent larval generations. Defoliation functions as a negative feedback mechanism acting on larval density. In 1989 local discoloration in the Upper Engadine valley was observed in the usual first focus. In 1990 and 1991, however, instead of the expected widespread defoliation damage, larval densities decreased drastically. Based on extensive field data collected from 1961 to 1991 on the development and the survival of the bud moth (Zeiraphera diniana Gn.) and the phenology of the host, Larix decidua L., this paper shows the effect of weather on survival in the egg stage and on the coincidence of larval hatching with the sprouting of the larch. It is shown that the winter and spring weather conditions in 1989–1991 were conducive to unusually high egg mortality. Since these conditions occurred in three successive generations, population growth was effectively reduced and the cycle collapsed prematurely. Thus the rather persistent cyclicity of the larch-larch bud-moth system was disturbed by weather conditions with a very low probability of occurrence, but due to the inherent high resilience of the system, the next population peak with visible defoliation is expected to occur 1996/1997, provided that the weather conditions return to the climatic standard.     

Numerical and temporal relationships in a three-level food chain

Summary Densities were examined in natural populations consisting of horsenettle (Solanum carolinense), larvae of a horsenettle-specific phytophagous moth (Frumenta nundinella), and a polyphagous parasitoid wasp (Scambus pterophori), that parasitized the moth. Herbivore and parasitoid densities were both positively associated with fruit density, but moth larval density was linear while parasitoid density increased parabolically; thus, optimum moth survivorship was at intermediate moth densities. There was no evidence of escape from herbivory in time by plant populations due to either mean flowering date or a spread of flowering in seasonal time. However, one plant population and one moth population apparently escaped in space due to isolated location. Although plant reproductive success is reduced whether or not moth larvae are parasitized, both presence of a numerical herbivore refuge and parasitoid attack at high fruit and moth densities would be expected to stabilize long-term temporal dynamics of this simple food chain.     

Density-dependent population dynamics in larvae of the dragonfly Pachydiplax longipennis: a field experiment

Summary Several features of dragonfly population biology suggest that population regulation occurs in the larval stage. This study was designed to determine if density-dependent interactions among larval odonates can affect survival, growth and emergence. First-instar larvae of the dragonfly Pachydiplax longipennis were raised in outdoor experimental ponds at initial densities of 38, 152, and 608 larvae · m^-2, under two levels of food availability. Food availability was supplemented in half the pools by volumetric addition of zooplankton every other day. Pools in the low food treatment did not receive the zooplankton supplement.There was a strong negative effect of density on the mean growth rate of survivors, which included both emerging tenerals and individuals overwintering in the larval stage. A higher proportion emerged from low density than high density pools. Metamorphs from high density populations were smaller and emerged slightly later than those from low density, but the absolute number of metamorphs did not differ significantly among density treatments. Food supplementation significantly increased the proportion of overwintering larvae. There were no significant food-by-density interactions, indicating that food and density acted independently on larval population dynamics. Density-dependent mechanisms can clearly contribute to odonate population regulation, especially by controlling the number of larvae which emerge and the average age at reproduction. Population-level responses to density may be a result of interference among larvae.     

Relative Importance of Environmental Stress and Herbivory in Reducing Litter Fall in a Semiarid Woodland

We examined the impact of soil stress (low water and nutrient availabilities) and two keystone insect herbivores on pinyon pine (Pinus edulis) needle litterfall. We compared trees growing on two distinct soil types: volcanic cinders, which exhibit pronounced water and nutrient limitation, and sandy-loam soils, which have higher water-storage capacity and nutrient availability. Using two long-term herbivore removal experiments (15 and 18 years, respectively), we also examined the effects of the pinyon needle scale (Matsucoccus acalyptus, which attacks juvenile trees) and the stem-boring moth (Dioryctria albovittella, which attacks mature trees) on pinyon litterfall. These herbivores reach high densities on cinder soils but are absent or occur at much lower levels on sandy-loam soils. Four years of litterfall measurements showed four major patterns. First, independent of herbivory, needle litterfall was 20% lower under trees on high-stress cinder soils than on sandy-loam soils. Second, in agreement with the negative impact of scales on tree growth (that is, a 30% decline in stem growth), trees with scale infestations had 25% lower litterfall rates than trees resistant to scale; however, 15 years of scale-insect removal did not significantly increase needle litterfall. This implies possible intrinsic differences in litter production between scale-resistant and scale-susceptible trees. Third, in contrast with significant negative effects of moth herbivory on tree growth (that is, a 27% decline in stem growth), moth herbivory had no effect on needle litterfall. This, along with increased stem density in moth-susceptible trees, may be evidence of compensatory production. Fourth, there were strong year by soil type and year by scale herbivory interactions, such that in some years the effect on litterfall can be obscured or reversed by some other factor. In summary, soil stress has a strong and predictable effect on needle litterfall, whereas the relationship between insect herbivory and needle litterfall is weaker and depends on the individual herbivore. These effects, however, are mediated by other environmental factors that have considerable annual variation.     

Food preference and performance of the larvae of a specialist herbivore: variation among and within host-plant populations

Specialist herbivores are suggested to be unaffected by or attracted to the defense compounds of their host-plants, and can even prefer higher levels of certain chemicals. Abrostola asclepiadis is a specialist herbivore whose larvae feed on the leaves of Vincetoxicum hirundinaria, which contains toxic alkaloids and is unpalatable to most generalist herbivores. The food choice, leaf consumption and growth of A. asclepiadis larvae were studied to determine whether there is variation among and within host-plant populations in their suitability for this specialist herbivore. There was significant variation in food preference and leaf consumption among host-plant populations, but no differences were found in larval growth and feeding on different host-plant populations. A. asclepiadis larvae preferred host-plant populations with higher alkaloid concentrations, but did not consume more leaf material from plants originating from such populations in a no-choice experiment. There was also some variation in food preference of larvae among host-plant individuals belonging to the same population, suggesting that there was variability in leaf chemistry also within populations. Such variation in larval preference among host-plant genotypes and populations may create potential for coevolutionary dynamics in a spatial mosaic.     

Interactions between Operophtera brumata and Tortrix viridana on oak: new evidence from time-series analysis

1. Yearly population estimates of the green oak leaf roller moth, Tortrix viridana (Lepidoptera: Tortricidae), and the winter moth, Operophtera brumata (Lepidoptera: Geometridae), were taken from the pedunculate oak, Quercus robur, from 1951 to 1966 in Wytham Woods, Oxfordshire, U.K. Larval mortality from parasitoids was recorded for both species over the same period. 2. Operophtera brumata population density fluctuated around a constant mean, whereas T. viridana population density exhibited a linear decline over the sampling period. Population counts were subjected to time-series analysis after the linear decline was removed from the T. viridana data by detrending. Multiple regression models were built so that variation in the per capita rate of increase of each population could be partitioned among (a) current and previous population density, (b) current and previous population density of the second moth species, and (c) rates of larval parasitism. 3. Multiple regression analysis suggested that variation in the per capita rate of increase of O. brumata could be explained by negative feedback from O. brumata density at T–2 (32%), a negative relationship with T. viridana density at T–1 (18%), and a positive relationship with parasitism suffered by T. viridana at T–1 (24%). 4. The T. viridana population time-series was dominated by a rapid feedback process such that per capita rate of increase at time T was negatively related to population density at time T–1, explaining 53% of variance in population growth rate. Per capita rate of increase was unrelated to larval parasitism or densities of O. brumata. 5. In light of previous life-table and experimental studies on O. brumata and T. viridana, the current analyses suggest that pupal predation (data not presented) and interspecific competition are significant determinants of O. brumata population growth rates in Wytham Woods. In contrast, T. viridana population growth rates appear to be dominated by rapid negative feedback consistent with intraspecific competition.     

Phenology and abundance in relation to climatic variation in a sub-arctic insect herbivore–mountain birch system

The two forest-defoliating geometrid moth species Operophtera brumata and Epirrita autumnata are known to exhibit different altitudinal distribution patterns in northern birch forests. One possible explanation for this is that altitudinal climatic variation differentially affects the performance of two species through mismatching larval and host plant phenology. We explored this hypothesis by investigating the relationship between larval phenology and leaf phenology of Betula pubescens, which is the main host plant of both moth species, along ten replicate altitudinal transects during two springs with contrasting climate in northern Norway. There was a distinct monotonous cline in host plant phenology with increasing altitude in both years of the study, but the development of the leaves were generally 14 days later in the first of the 2 years due to cold spring weather. We found that larval development of both species closely tracked host plant leaf phenology independent of altitude and year. However, at the time of sampling, E. autumnata was approximately one instar ahead of O. brumata at all altitudes, probably reflecting that E. autumnata has faster early instar growth than O. brumata. The abundance of O. brumata was lowest at the altitudinal forest-line, while E. autumnata was lowest near sea level. Our results do not indicate that the altitudinal distribution patterns of the two moth species is due to any phenological mismatch between larval and host plant phenology. We suggest rather that natural enemies at low altitudes limit larval survival and thus abundance of E. autumnata, while an early onset of winter at the forest limit reduces survival of late eclosing adults of O. brumata.     

Response to long- and short-term salinity in populations of the C4 nonhalophyte Andropogon glomeratus Walter B.S.P.

Summary This research was undertaken to investigate differences in salt tolerance under conditions in which salinity is increased gradually and maintained for long periods or increased rapidly and maintained for shorter periods. The responses of populations of a C₄ nonhalophytic grass, Andropogon glomeratus, to long- and short-term salinity were measured under controlled environment conditions. Additionally, plants from a salt marsh population and an inland population were transplanted into a salt marsh and their survival compared. The relative growth reductions in the salt marsh and the inland populations under long-term salinity were similar. Survival of seedlings of 4 populations inundated with full-strength seawater over a relatively short period indicated differential capacities to tolerate soil salinities imposed in a manner similar to tidal inundation in a salt marsh. The greater survival of plants from the marsh population transplanted into the salt marsh further indicated genetic differentiation between the populations. These results indicate that genetic differentiation to salt tolerance in A. glomeratus is better reflected by survival after shortterm salinity events, rather than growth inhibition due to long-term salinity imposed gradually.     

Reduced parasitism of a leaf-mining moth on trees with high infection frequencies of an endophytic fungus

We investigated relationships between endophytic fungi and a leaf-mining moth, Phyllonorycter sp., along an elevational gradient from 2255 to 2895 m. The fungi and moth larvae inhabit leaves of Quercus gambelii. Fungal frequencies and larval densities varied with elevation. However, larval densities were not associated with the frequencies of infection by endophytic fungi. Survival of larvae was positively associated with the most dominant fungus, Gnomonia cerastis, owing to reduced parasitism of moth larvae on trees with high frequencies of Gnomonia.     

The effect of a predatory leech,Nephelopsis obscura,on mortality,growth, and production of chironomid larvae in a small pond

Summary The effect of a predatory leech, Nephelopsis obscura, on survivorship, growth, and production of chronomid larvae was studied by enclosure experiments carried out in a small pond. The prey population was composed almost entirely of the tubiculous, microphagous chironomid larvae, Chironomus riparius and Glyptotendipes paripes. Nephelopsis significantly reduced chironomid survivorship within the enclosures, and accounted for most of the measured mortality of fourth instar larvae. The cropping by Nephelopsis was not significantly biased toward either prey species. In long-term experiments (66 d) chironomid biomass in enclosures without leeches reached much higher levels than in enclosures containing Nephelopsis. This increase in biomass was due to growth of surviving larvae, rather than recruitment, since emergence and oviposition were not going on during the course of the experiments. The enhanced survivorship of larvae within leech-free enclosures was eventually accompanied by reduced growth and specific production (daily production/biomass) for C. riparius, which made up about 90% of the larval population. Growth and specific production of G. paripes (10% of larval population) was not affected. Short-term experiments (25 d) involving manipulation of densities and species ratio (9:1 CR:GP and 1:9 CR:GP) of larvae revealed that growth of the majority species was strongly influenced by larval density, whereas growth of the minority species was not. The same pattern was observed both in the presence and in the absence of Nephelopsis and was a result of differences in resource utilization between the two species. In the shortterm experiments, growth rates estimated for larvae exposed to leeches were significantly less than those for larvae in leech-free enclosures. This could be due either to size-biased consumption of larvae by Nephelopsis, or possibly a disturbance factor leading to reduced larval food intake and/or increased metabolic costs.     

Exploration into a mechanism of transgenerational transmission of nucleopolyhedrovirus in Lymantria dispar L. in Western Siberia

The mechanisms of nucleopolyhedrovirus (NPV)transmission through gypsy moth (Lymantriadispar L., Lepidoptera: Lymantriidae)populations in Western Siberia wereinvestigated. Field and laboratory studies werecarried out during 1992–1998 in a gypsy mothoutbreak in the Novosibirsk Area (WesternSiberia), Russia. The damaged trees were primarilytwo species of birch, Betula pendula Roth. and B. pubescensEhrh. Investigations were performed at threestages of the outbreak: population increase(1992–1993), population peak (1994–1995) andpopulation decline (1996–1998). Experiments onactivation of occult virus by different stressfactors were carried out in order to clarifythe mechanism of transgenerational transmissionof NPV. In both laboratory and fieldexperiments NPV-caused mortality levels werelow; most insects did not acquire a lethal doseof virus. Occult virus can provide an importantroute of transgenerational NPV transmission,particularly in Western Siberia where gypsymoths migrate by female flight and can moveaway from trees contaminated by virus fromprevious larval infections.     

Effect of habitat complexity on biological control by the red imported fire ant (Hymenoptera: Formicidae) in collards

A potentially important and understudied biological control agent in US agroecosystems is the red imported fire ant, Solenopsis invicta Buren. Red imported fire ants may be particularly important biological control agents because we can manipulate their abundance with changes in habitat complexity. The effect of habitat complexity on biological control by fire ants was determined using plots of collards intercropped with white clover (complex habitat) and simple collard monocrops. The most economically significant pests of collards are larvae of the diamondback moth (DBM), Plutella xylostella (L.). Predation of DBM larvae by fire ants was more rapid and efficient in the intercrop than the monocrop. Red imported fire ants were 23% less abundant in the intercrop than the monocrop, however, suggesting that fire ants had a greater per capita effect on DBM survival in the complex habitat. Red imported fire ant predation of DBM larvae was significantly affected by larval density. Red imported fire ants also reduced the survival of leaf beetles, another economically significant pest taxa, by 45%. Furthermore, collard leaf damage tended to be inversely related to fire ant density and fire ants were more effective at reducing crop damage in the complex intercrop. Our study indicates the ability of red imported fire ants to be effective biological control agents and suggests that increasing habitat complexity can enhance red imported fire ant efficacy and herbivore control.     

Delayed inducible resistance against a leaf-chewing insect in four deciduous tree species

Summary Both mechanical damage to mountain birch foliage and rearing of moth larvae on the trees reduced the growth of Epirrita autumnata larvae reared on these trees in the following year. The effects of physical damage and some other cues from insects were additive. On bird cherry the performance of Epirrita larvae was equal on untreated trees and on trees artificially defoliated in the previous year, but larval growth was reduced on previously insect-damaged branches. With mountain ash just physical damage per se reduced the performance of Epirrita larvae. On Salix phylicifolia there were no significant differences in the growth or survival of Epirrita on untreated control bushes and on bushes with partial larval damage during the previous year. Among untreated control trees the growth and survivorship of Epirrita were higher on fast-growing willow and bird cherry than on the slow-growing mountain birch. Mountain birch and mountain ash, the two deciduous tree species adapted to nutrient-poor soils, showed delayed inducible resistance triggered by defoliation (artificial or insect-made). This supports the hypothesis that delayed inducible resistance may be a passive response due to nutrient-stress caused by defoliation. On the other hand, the additional increase in the resistance of mountain birch triggered by specific cues from insects suggests that this response may be an evolved defense against leaf-eating insects.     

Does prescribed burning mean a threat to the rare satyrine butterfly Hipparchia fagi? Larval-habitat preferences give the answer

The ecological effects of fire management, especially regarding arthropods are poorly investigated. Burning in winter was assumed to pose a threat to butterfly species hibernating as larvae. To assess the impact of prescribed burning on population viability, we analysed larval-habitat preferences of the highly endangered, xero-thermophilous butterfly Hipparchia fagi in vineyards of the Kaiserstuhl region (southern Germany). Microhabitat preference analyses for mature larvae and egg-laying females revealed a preference of H. fagi for Bromus erectus-dominated communities with sparse vegetation coverage and a distinct tuft growth of the host plant B. erectus on microclimatically benefited slopes. We explain the preference of B. erectus by a preference of vegetation structure. The grass tufts offer a suitable climatically buffered living space for larvae. Egg deposition took place on dry substrate at positions of high solar radiation, thus adapted to hot and dry microclimate. As the larval habitat was sparsely vegetated as well as generally legally protected, fire management was not applicable and therefore not affecting the populations. We think it is conceivable that H. fagi, occurring here at its northern range limit, might expand its larval habitat into denser, combustible B. erectus stands in the course of global warming. A change in habitat preferences would necessitate a re-evaluation of management options.     

Experimental simulations of climate change induced mismatch in oak and larval development rates impact indicators of fitness in a declining woodland moth

Consequences of climate change-driven shifts in the relative timing of spring activities of interacting species are insufficiently understood, especially for insects. We use a controlled experiment which simulates a trophic mismatch scenario in which lepidopteran larvae predominately feed on older leaves due to foliage developing faster than larvae growth rates. As a case study our experiment uses Orthosia cerasi, which is a widespread but declining woodland moth whose UK declines appear to be driven by warming temperatures. In the control experiment larvae are fed young oak Quercus robur leaves (bud burst stages six and seven), whilst in the treatment newly emerged larvae are fed young leaves but then gradually transition to feed on older leaves (post bud burst stage seven). We assess impacts on duration of the larval stage, pupal size and overwintering duration and survival. Larvae in the phenological mismatch treatment had a longer larval period, and smaller and lighter pupae. Larval diet did not carry over to influence emergence dates as earlier pupation of control larvae was balanced by an equivalent increase in the duration of the pupal stage. Increased time spent as larvae could increase predation rates from avian predators, whilst slowing the seasonal decline in food availability for those bird species. Reduced pupal size and weight are indicators of lower fecundity in emerging adults. Notably, we find that adults emerging from the mismatch treatment exhibited greater rates of abnormal vestigial wing development, which is likely to further reduce fitness. Trophic mismatches in which caterpillars have reduced availability of young leaves may thus contribute to the population declines observed in many woodland moth species due to increased mortality at larval stages, and adverse effects of early life conditions that reduce the reproductive success of emerging adults.     

Conserving the New Forest burnet moth (Zygaena viciae ([Denis and Schiffermueller])) in Scotland; responses to grazing reduction and consequent vegetation changes

Zygaena viciae, the New Forest burnet moth, has only one population in Britain, in western Scotland. Here it was discovered in1963 and its population sustained itself, before declining seriously from 1980 to 1990. A survey in 1990 discovered at most 20 adult moths and it was clear that the site had become seriously over-grazed. A fence was erected to exclude sheep, with variable success until 1996, since when it has remained effective. Vegetation speedily changed from 1990 onwards, including re-establishment and spread of the main larval foodplant, Lathyrus pratensis. The moth population remained low until 1997, since which time it has dramatically increased, reaching an estimated 8500–10,200 in 2003. However, with only one site the moth remains threatened and establishment on new sites is now a priority.     

How do food quality and larval crowding affect performance of the autumnal moth,Epirrita autumnata?

Outbreak densities of autumnal moth, Epirrita autumnata (Borkhausen) (Lepidoptera: Geometridae), lead to high larval crowding. Phenotypic responses of E. autumnata to larval crowding and to food quality were studied by measuring growth and consumption as well as pupal weight and fecundity. Crowding may trigger increased consumption and faster development to avoid impending food shortage on good quality food. This is suggested by the result that on a good‐quality diet, the growth of crowded larvae was better than that of solitary larvae, though they did not consume more food than solitary larvae. Crowded larvae also completed the last instar earlier than solitary larvae. The fecundity of crowded autumnal moths was not lower than the fecundity of solitarily grown autumnal moths. This may provide conditions for extra rapid population build‐up of E. autumnata. During the population increase phase the crowding effect may facilitate larval performance; however, at peak density the crowding starts to have negative effects on the performance of larvae. On a poor‐quality diet, the performance of crowded and solitary larvae did not differ. The growth of larvae was better on a good‐quality diet than on a poor‐quality diet, due to higher efficiency in food utilization. Larvae feeding on low‐quality diet did not prolong their development time, but pupated at smaller size; this resulted in lower fecundity. A decrease in food quality can be seen as a cue of oncoming food shortage and resource depletion; it may be advantageous to pupate at a smaller size and ensure survival till reproduction, rather than risk prolonging development to achieve larger size and higher fecundity.