Larval parasitoids and pathogens of the fall armyworm in honduran maize

Parasitism by natural enemies of all larval instars of the fall armyworm (FAW),Spodoptera frugiperda, was quantified in 5 Honduran maize fields during the 1984 growing season. Overall, 42.0% of the host larvae were killed by the natural enemy complex. The braconid parasitoidChelonus insularis was the most common natural enemy accounting for 36.8% of the complex and causing 15.5% mortality of FAW larvae. Other important natural enemies included the nematodeHexamermis sp., the tachinidLespesia sp. and the imperfect entomopathogenic fungusNomuraea rileyi. Distinct ranges of larval instars were attacked by several natural enemy species.Chelonus insularis was an important FAW natural enemy at all larval densities observed. Florida Agricultural Experiment Station Journal Series No. 8650 and MIPH-EAP No. 146.     

Leaf flushing phenology and herbivory in a tropical dry deciduous forest,southern India

Patterns of leaf-flushing phenology of trees in relation to insect herbivore damage were studied at two sites in a seasonal tropical dry forest in Mudumalai, southern India, from April 1988 to August 1990. At both sites the trees began to flush leaves during the dry season, reaching a peak leaf-flushing phase before the onset of rains. Herbivorous insects emerged with the rains and attained a peak biomass during the wet months. Trees that flushed leaves later in the season suffered significantly higher damage by insects compared to those that flushed early or in synchrony during the peak flushing phase. Species whose leaves were endowed with physical defenses such as waxes suffered less damage than those not possessing such defenses. There was a positive association between the abundance of a species and leaf damage levels. These observations indicate that herbivory may have played a major role in moulding leaf flushing phenology in trees of the seasonal tropics.     

The phenological window of opportunity for early-season birch sawflies

Abstract 1. The phenological window of opportunity hypothesis posits that an insect herbivore cannot survive outside a specific leaf age interval of its host plant.
2. Previous studies have shown that mountain birch displays extensive seasonal changes in leaf biochemical characteristics. Young, expanding leaves are full of water and amino acids but they also contain large amounts of protein-precipitating gallotannins as well as flavonoid-glycosides. Concentrations of these compounds decline during leaf growth whereas concentrations of sugars and proanthocyanidins and leaf toughness increase.
3. Adult birch sawflies hatch in early summer and oviposit on growing leaves, which compels larvae to feed on leaves that are already well developed. The purpose of this study was to test whether leaves that were younger and biochemically different from the leaves available under natural conditions are within the phenological window of opportunity for the larvae of two early-season birch sawfly species, Amauronematus amplus Konow and Pristiphora alpestris (Konow).
4. Amauronematus amplus larvae survived better and developed faster, and P. alpestris larvae developed faster and became bigger, on atypically young leaves compared with larvae reared on leaves encountered normally. Therefore, these species can exploit the putative nutritional superiority of very young leaves, which probably outweighs the potential impact of the new set of secondary metabolites offered to them. In conclusion, young leaves that are consumed rarely by sawfly larvae are within their phenological window of opportunity, even though the timing of sawfly life cycles constrains their utilisation.     

Plant phenology-mediated indirect effects: The gall midge opens the phenological window wider for a leaf beetle

We examined whether larvae of the gall midge Rabdophaga rigidae (Diptera: Cecidomyiidae) can modify the seasonal dynamics of the density of a leaf beetle, Plagiodera versicolora (Coleoptera: Chrysomelidae), by modifying the leaf flushing phenology of its host willow species, Salix serissaefolia and Salix eriocarpa (Salicaceae). To test this, we conducted field observations and a laboratory experiment. The field observations demonstrated that the leaf flushing phenology of the willows and the seasonal dynamics of the beetle density differed between shoots with stem galls and shoots without them. On galled shoots of both willow species, secondary shoot growth and secondary leaf production were promoted; consequently, leaf production showed a bimodal pattern and leaf production periods were 1 to 2 months longer than on non‐galled shoots. The adult beetle density on galled shoots was thus enhanced late in the season, and was found to change seasonally, synchronizing with the production of new leaves on the host willow species. From the results of our laboratory experiment, we attributed this synchrony between adult beetle density and willow leaf flush to beetles’ preference to eat new leaves rather than old. Indeed, beetles consumed five times more of the young leaves when they were fed both young and old leaves. These results indicate that stem galls indirectly enhance the adult beetle density by enhancing food quality and quantity late in the beetle‐feeding season. We therefore conclude that midge galls widen the phenological window for leaf beetles by extending the willows’ leaf flush periods.     

Performance of a spring-feeding moth in relation to time of oviposition and bud-burst phenology of different host species

Abstract. 1. As a spring-feeding moth committed to immature foliage, the autumnal moth Epirrita autumnata (Lepidoptera, Geometridae) must have egg hatch synchronised with the bud-burst of its host plants. Due to large individual variation in the length of the pupal period, however, E. autumnata populations exhibit a prolonged period of flight and oviposition. Because the timing of oviposition in autumn is associated with the timing of egg hatch in the following spring, the time window for egg hatch expands and more potential hosts may become attainable. This suggestion was evaluated under field conditions by rearing E. autumnata eggs and larvae on four different hosts.
2. The performance of E. autumnata was measured by using estimates for fecundity (pupal mass) as well as survivorship of eggs and larvae. Based on the availability of foliage and phenological synchrony between larval and leaf development, early-laid eggs and the larvae originating from them were predicted to perform better on the hosts that have early-flushing leaves. On the late-flushing hosts, the larvae that hatched later were predicted to perform better than the larvae that hatched earlier. Half of the trials were exposed to predators and parasitoids, while the rest were conducted inside mesh-bags preventing larval dispersal and mortality due to natural enemies.
3. The results of the experiment did not support the simple predictions. In particular, host-plant quality and natural enemies appeared to operate discordantly between early- and late-laid eggs. Larvae from the late-laid eggs had rapid development during the larval stages and pupated at the same time and with the same pupal mass as the larvae hatched from the early-laid eggs.
4. The results indicate an occurrence of several, unknown selective forces in E. autumnata populations maintaining variation in the length of the pupal period, timing of oviposition, and timing of egg hatch.     

Influence of leaf trichome density on the efficiency of two polyphagous insect predators

Plant characteristics, such as leaf structure or hairiness, are important for the movement and attachment of insects. It has been suggested that increased trichome density on new Salix cinerea L. (Salicaceae) leaves, produced after grazing by the willow leaf beetle Phratora vulgatissima L. (Coleoptera: Chrysomelidae), function as an inducible defence against the beetle and especially its larvae. Here we studied whether and how two of the main natural enemies of P. vulgatissima, viz., Anthocoris nemorum L. (Heteroptera: Anthocoridae) and Ortothylus marginalis L. (Heteroptera: Miridae), were influenced by trichome density on S. cinerea leaves. The effect of trichome density on these two predators was studied on plants with different trichome densities, comparing natural enemy efficiency, measured as number of P. vulgatissima eggs consumed or larvae missing and/or killed. To obtain different trichome densities, cuttings of several different clones of S. cinerea were used. In the experiment using eggs as prey, an increase in trichome density was, in addition, induced through leaf beetle defoliation on half of the plants of each willow clone. Furthermore, a field study was performed to investigate whether trichome density was correlated with natural enemy abundance. The results indicate that neither the efficiency of these two natural enemies in the greenhouse, nor their abundance in the field was influenced by trichome density. A well‐known behavioural difference between the two predator species could probably account for the higher disappearance of larvae after exposure to the more active predator. These findings are relevant for the development of pest management programs, not least because the enemies are polyphagous predators. It is concluded that an induced increase in leaf hairiness in willows in response to leaf beetle grazing could be a plant resistance trait worthy of further study in this system, because no negative effects on the main natural enemies were observed.     

Timing is everything? Phenological synchrony and population variability in leaf‐chewing herbivores of Quercus

Abstract.  1. Specialization on ephemeral resources (e.g. new leaves) should produce large annual variation in herbivore population size when the timing of availability of those resources is unpredictable. Despite considerable evidence for impacts of synchrony with budburst on survival of larval Lepidoptera, previous studies of adult Geometridae and Noctuidae found no correlations between insect phenology and population variability.
2. We surveyed larval Lepidoptera feeding on Quercus alba and Q. velutina in Missouri from 1993 to 2003 and examined population variability, measured as the coefficient of variation of population density (CV), in a subset of abundant species. We compared CV values among species whose larvae feed only in spring, early summer, mid-summer, late summer, or all season. We predicted that univoltine species whose larvae eclose and complete development in spring during leaf expansion would have higher variability than species feeding later in the season, having multiple generations, or having longer development times.
3. As predicted and consistent with hypotheses, spring-feeding species had CV values 32% higher than species feeding in summer months. Coefficients of variation were also 34% higher in leaf-rolling and mining guilds compared with free-feeders, suggesting that mobile species may compensate for asynchrony with budburst by dispersing to higher quality plants or plant parts. Multivoltine species, however, did not differ from univoltine species in population variability.
4. Our results suggest that asynchrony with plant phenology and factors that might exacerbate it, such as climate change, will have the largest impacts on the dynamics of spring-feeding Lepidoptera, particularly species with limited mobility.     

Does low nutritional quality act as a plant defence? An experimental test of the slow‐growth,high‐mortality hypothesis

Abstract.  1. The slow-growth, high-mortality hypothesis was experimentally tested in this study by investigating the effects of plant quality and natural enemies on leaf-miner growth, performance, and survivorship. Two leaf miners ( Acrocercops albinatella and Brachys tesselatus ) occurring on the turkey oak Quercus laevis were studied using a factorial design that manipulated predation/parasitism pressure and plant nutritional quality.
2. Forty trees were randomly divided into four treatments: (1) control plants (nutrients and natural enemies unaltered); (2) nutrients added, natural enemies unaltered; (3) nutrients unaltered, natural enemies reduced; and (4) nutrients added and natural enemies reduced. Water content, leaf toughness, tannin concentration, and foliar nitrogen were quantified monthly for each plant, and mine growth and survivorship were assessed by tracing mines on a 2–3-day interval and by following the fates of 50 mines per species per treatment combination.
3. Fertilised plants exhibited significantly higher amounts of nitrogen, but no significant differences among treatments were observed for water content, leaf toughness, and tannin concentration. These results only partially support the slow-growth, high-mortality hypothesis, as mines were significantly smaller and developed faster on fertilised plants, but neither fertilisation nor natural enemy exclusion significantly affected mine survivorship or mortality caused by natural enemies.     

Opposing effects of spring defoliation on late season oak caterpillars

ABSTRACT. 1. The pedunculate oak, Quercus rohur L., suffers high annual levels of spring defoliation in Wytham Woods. near Oxford.
2. This spring defoliation affects late season caterpillars through a variety of damage-induced changes in the leaves.
3. Diurnea fagella (D. & S.), one of the commonest late season caterpillars, shows reduced larval survival and pupal weight on regrowth foliage when compared to undamaged primary foliage.
4. D. fagella also suffer higher larval mortality on naturally damaged primary foliage than they do on undamaged foliage.
5. Despite this, the three commonest late season caterpillar species are more abundant on damaged trees than undamaged ones. and their distributions are biased towards damaged leaves within the canopy.
6. Other factors that may be more important than leaf damage in determining the distribution and abundance of late season caterpillars are discussed. D.fagella larvae spin their larval refuges more rapidly on damaged than undamaged foliage, and this may reduce mortality by natural enemies, or ameliorate adverse effects of weather.     

Biology,natural history and temporal fluctuation of the geometrid Oospila pallidaria associated with host plant phenology

We investigated the temporal fluctuation of the geometrid moth Oospila pallidaria and related it to the leaf flush season of its host plant, Mimosa setosa, and to environmental factors. Aspects of the natural history of adults are also described. Mimosa setosa produced leaves all year round, but abundance of O. pallidaria larvae in the field was seasonal. The abundance of immatures showed a tendency to decrease with rainfall and temperature. Our study is pioneering in that it provides insights into factors related to the occurrence of O. pallidaria in natural areas. In addition, the biology of immatures is provided.     

Changing dynamics of the pine beauty moth (Panolis flammea) in Britain: the loss of enemy free space?

1 The pine beauty moth Panolis flammea has two main host plants in Britain: Pinus sylvestris (Scots pine), which is the ancestral food plant where the insect is never abundant enough to cause tree mortality, and Pinus contorta (lodgepole pine), an introduced host tree that has experienced periodic widespread tree mortality due to this pest.
2 We review the recent literature, published mostly after the year 2000, regarding the impact of natural enemies on the population dynamics of P. flammea in Britain.
3 The natural enemies of P. flammea are more diverse and abundant in Scots pine habitat than in lodgepole pine habitat and some of them show differential selection for P. flammea larvae in Scots pine habitat over those located in lodgepole pine habitat.
4 It is concluded that the difference in the population dynamics of this insect in the two different habitats was probably the result of the P. flammea finding enemy-free space in lodgepole pine habitat.
5 Recent evidence on the diversity and impact of natural enemies on lodgepole pine has demonstrated that they currently have a much more significant impact on this pest than they did in the 1970s and 1980s, when outbreaks were frequent.     

Understanding gregariousness in a larval Lepidopteran: the roles of host plant, predation, and microclimate

Abstract.  1. Many moth and butterfly larvae are gregarious early in development, but become solitary in late instars. This ontogenetic variation in behaviour is probably the result of temporal changes in the costs and benefits associated with gregariousness. This study provides observational and experimental evidence that, in one particular moth species, a series of different ecological factors influence larval behaviour at different times during development.
2. Field observations show that young caterpillars of the limocodid Doratifera casta form large aggregations while foraging, but that mature larvae are largely solitary.
3. A field experiment revealed that individual first to third instar larvae in larger groups develop more rapidly, but that group size had no detectable influence on survival. The developmental advantage associated with gregariousness is affected by host plant species, but not by predator exclusion, suggesting that group living in these cryptic early instar larvae promotes feeding facilitation, but does not provide individuals with protection from natural enemies.
4. Laboratory experiments revealed that aposematic fourth instar caterpillars in large groups were less likely to be attacked by a generalist insect predator than those in small groups.
5. Field observations provided no evidence that group living affects body temperature, suggesting that microclimatic factors do not favour gregariousness in this species.
6. It is concluded that gregariousness in D. casta confers at least two different advantages on larvae at different stages early in development, but that these advantages disappear, or are outweighed by costs associated with intraspecific competition, in final instars.     

Testing the importance of nest concealment: does timing matter?

ABSTRACT.   Nest concealment by vegetation is considered an important factor affecting predation rates for many passerines and, therefore, is frequently measured in studies examining nest predation. However, the time when concealment measurements are made may affect the results of such studies, particularly in highly seasonal ecosystems where characteristics of the vegetation later in the breeding period may differ considerably from those at the time of nest-site selection. We used artificial nests baited with quail ( Coturnix sp.) eggs in a highly seasonal tropical dry forest in Jalisco, western Mexico, to test the effects of seasonal change in concealment on nest predation. We placed 40 open-cup, artificial nests in shrubs at the end of the dry season and again at the beginning of the rainy season in 2007, and monitored the fate of the nests and the degree of concealment by vegetation during both periods. Nest concealment was significantly greater during the wet season than during the dry season. The percentage of nests predated was marginally higher during the dry (100%) than the wet (72.5%) season, and daily nest survival was lower during the dry than the wet season. Our results suggest that, in highly seasonal environments such as tropical dry forests, delayed measurement of nest concealment after nest completion rather than during nesting may constitute a significant source of error.     

Early spring defoliation,secondary leaf flush,and leafminer outbreaks on American holly

Summary Adults of Phytomyza ilicicola (Diptera: Agromyzidae), a univoltine specialist leafminer, emerge in close synchrony with leaf flush of American holly and feed on and oviposit in soft, partially expanded leaves. Early spring defoliation, such as commonly results from freezing injury to young shoots, is followed several weeks later by a second flush of young leaves from lateral buds. We simulated this phenomenon by manually defoliating whole small trees and individual shoots of large trees to test the hypothesis that freezing injury can encourage leafminer outbreaks by inducing an abundance of soft, protein rich young leaves late in the adult activity period, when availability of vulnerable leaves becomes limited. Defoliation of small trees one or two weeks after bud break resulted in six- to 13-fold increases in the incidence of feeding punctures and larval mines on second flush leaves as compared with densities on original young leaves of control trees. Similarly, we induced significant increases in feeding punctures and larval mines on second flush leaves of individual defoliated shoots, although leaves that did not open until after the flight period escaped this injury. These observations underscore the capability of adult female P. ilicicola to locate and exploit a small number of phenologically available leaves among many hundreds of older leaves on the same tree. By altering the phenology of leaf flush, certain kinds of environmental stress may predispose perennial plants to outbreaks of early season folivores that restrict their feeding or oviposition to very young leaves.     

Host tree architecture mediates the effect of predators on herbivore survival

Abstract.  1. Vegetation structural complexity is an important factor influencing ecological interactions between different trophic levels. In order to investigate relationships between the architecture of trees, the presence of arthropod predators, and survival and parasitism of the autumnal moth Epirrita autumnata Borkhausen, two sets of experiments were conducted.
2. In one experiment, the architectural complexity of mountain birch was manipulated to separate the effects of plant structure and age. In the other experiment the trees were left intact, but chosen to represent varying degrees of natural complexity. Young autumnal moth larvae were placed on the trees and their survival was monitored during the larval period.
3. The larvae survived longer in more complex trees if predation by ants was prevented with a glue ring, whereas in control trees smaller canopy size improved survival times in one experiment. The density of ants observed in the trees was not affected by canopy size but spider density was higher on smaller trees. The effect of canopy structure on larval parasitism was weak; larger canopy size decreased parasitism only in one year. Until the fourth instar the larvae travelled shorter distances in trees with reduced branchiness than in trees with reduced foliage or control treatments. Canopy structure manipulation by pruning did not alter the quality of leaves as food for larvae.
4. The effect of canopy structure on herbivore survival may depend on natural enemy abundance and foraging strategy. In complex canopies herbivores are probably better able to escape predation by ambushing spiders but not by actively searching ants.     

Opposing survivorship and fecundity effects of host phenology on the gall-forming aphid Hormaphis hamamelidis

Abstract  1. This study examined the synchronisation and roles of host and insect phenology for fundatrices of the gall-forming aphid Hormaphis hamamelidis , which forms galls on the leaves of the understorey tree Hamamelis virginiana .
2. Host and insect phenologies varied widely among years and individual host trees.
3. Most winter eggs (≈ 85%) hatched before budbreak and galls could be initiated but in 2 years there was no inherent risk for earlier hatching fundatrices in terms of greater mortality. Egg hatch was not synchronised tightly with host trees but instead seemed to occur well in advance of budbreak as a conservative, bet-hedging strategy to ensure the opportunity to form galls.
4. Fundatrices that formed galls later had higher fecundity, contrary to typical patterns for insects that feed on the first flush of leaves in temperate forests, where later herbivores have lower fecundity. Also, unlike many other systems, trees that had later budbreak had greater densities of galls.     

Can life‐history and defence traits predict the population dynamics and natural enemy responses of insect herbivores?

Abstract.  1. Life-history differences between herbivorous insects with eruptive and latent population dynamics are potentially useful for predicting population size variability. An association has also been demonstrated between herbivorous insect defence traits and the responses of various natural enemies.
2. Here predictions of population dynamics and natural enemy responses based on life-history and defence traits are tested using Gonometa postica Walker and G. rufobrunnea Aurivillius, two Southern Hemisphere Macrolepidoptera (Lasiocampidae) species. The temporal and spatial variation in pupal abundance and patterns of pupal parasitism and predation for both species are described and quantified for the first time.
3. Eleven sites were sampled over four generations across the region where both species have historically reached high population densities. Although there was evidence suggesting that population synchrony is driven by weather patterns, site-specific environmental differences contributed to the observed population variability. This study is the first to quantify the extent of population size variability of a species with an intermediate position on the eruptive–latent population dynamic gradient, where data on insect population dynamics is scarce.
4. Support for the life-history–population dynamic relationship was found, as intermediate population size variability for these species was observed. Larval and pupal defence traits, however, were poor and inconsistent predictors of mortality rate. Pupal cocoon structure differences, previously documented for these Gonometa species, may in fact explain the interspecific differences in natural enemy responses found.
5. Predicted population dynamics and natural enemy responses may, however, be overridden by ecological conditions. Nevertheless, life-history and defence traits provide a useful basis for predicting population dynamics of poorly studied species.     

Evidence for host-associated races in a gall-forming midge: trade-offs in potential fecundity

Abstract. 1. The gall midge, Asphondylia borrichiae , attacks the terminals of three plants in the aster family: Borrichia frutescens , Iva frutescens , and I. imbricata .
2. In the field, Borrichia suffers the highest rate of galling and I. imbricata the lowest. Common garden experiments also revealed highly significant differences in the attack rates of the three host species by Asphondylia . However, these differences depended on the source of the attacking midge population.
3. Midges collected from Borrichia and I. frutescens attacked the host in which they developed almost exclusively, whereas those from I. imbricata attacked both Iva spp.
4. Each Asphondylia larva develops in its own chamber within a gall. Borrichia galls were 2.5–3.5-fold less crowded than those from I. imbricata and I. frutescens , respectively. Consequently, midges that developed in Borrichia were significantly larger and eclosed with 30–40% more eggs than those that developed in I. imbricata and I. frutescens , respectively.
5. Although performance of Asphondylia larvae was lowest on the two Iva spp., especially I. frutescens , these hosts may provide an escape from natural enemies as a trade-off for reduced offspring performance.
6. Differences in host-plant species phenology may reduce gene flow among host-associated populations of Asphondylia , thereby favouring the formation of races at the level of plant genus.     

东北地区大豆田天敌昆虫优势种类分析

2005~2007年,对中国哈尔滨地区大豆田天敌昆虫开展系统调查。共发现5目、12科、33种天敌昆虫。龟纹瓢虫Propylaea japonica(Thunberg)、异色瓢虫Harmonia axyridis(Pallas)、隐翅甲Ischnosoma sp.成虫和花蝽Oriussp.为大豆田优势种天敌昆虫。其中,隐翅甲Ischnosomasp.成虫于6月上旬至下旬田间发生。花蝽Oriussp.成虫6月下旬至8月下旬发生,异色瓢虫成虫6月下旬至9月下旬发生。龟纹瓢虫幼虫、异色瓢虫幼虫和花蝽Oriussp.若虫多于6月下旬至8月下旬,即大豆生育中期发生;龟纹瓢虫成虫田间发生期较长,大豆生育期中6月中旬至9月下旬均有发生。     

Differences in leaf phenology between juvenile and adult individuals of two tree species in a seasonally dry tropical woodland

In temperate forests, juvenile trees anticipate leaf phenology compared to adults, thus avoiding shading and herbivory. This is also expected to occur in seasonal tropical forests due to intense herbivory and shading during the rainy season; however, the anticipation of leaf phenology by juveniles in seasonal tropical forests has yet to be demonstrated. Stem‐succulent species are expected to be prone to juvenile phenological anticipation because these species are able to use water stored in their stems for leaf flushing in the dry season. We investigated this hypothesis by comparing leaf phenology (bud break, leaf expansion) of juveniles and adults of two species with contrasting wood densities in the transition between dry and rainy seasons in a tropical dry woodland. We also investigated the level of light limitation that juveniles experience in the rainy season. Both species exhibited bud break during the dry season, but only expanded their leaves with the occurrence of the first rains. In general, the stem‐succulent species had a more precocious bud break; however, anticipation by juveniles occurred only in the species with more dense wood. Canopy openness was lower than in temperate deciduous forests, but the fact that the full expansion of leaves occurred only with rainfall indicates that bud break in anticipation of canopy closure contributes only to keeping leaf photosynthetic balance from going negative, and not to higher carbon gain. The importance of anticipated budding for escaping herbivory remains an alternative explanation in need of investigation.