Life history and behavioural traits of Mnesampela privata that exacerbate population responses to eucalypt plantations: Comparisons with Australian and outbreak species of forest geometrid from the Northern Hemisphere

The larvae of the native Australian moth Mnesampela privata (Geometridae) sometimes defoliate plantation eucalypts, causing concern to industry. Typically, populations of M. privata maintain innocuous numbers in native forests, but outbreak populations can occur in plantations. Certain of the life history and behavioural traits of M. privata exacerbate population responses in simplified systems. Specifically, M. privata exhibits indiscriminate oviposition behaviour, for example, females oviposit upon leaves where conspecific egg clutches are already present. Combined with large egg clutches, this trait can lead to heavy exploitation of natal hosts. Complete defoliation of the natal tree necessitates that larvae disperse to undamaged hosts. In native forests, dispersing larvae would have a lower probability of locating a new host of suitable species and phenotype than they would in a plantation. Larval tolerance of a wide variety of species of Eucalyptus facilitates utilization of non‐natal eucalypts. A lower rate of encounter between larvae and natural enemies is achieved by means of leaf shelters and autumnal seasonal activity. Good dispersion by gravid females from plantations with large moth populations ensures that nearby plantations will be located. Given these characteristics, continued outbreaks of M. privata are likely in monospecific eucalypt plantations, especially when management practices reduce populations of natural enemies and overall vegetational complexity. However, because outbreaks only arise in plantations, population fluctuations of M. privata cannot be considered ‘self‐driven’. Hence, this moth is not a true eruptive species in the manner of other geometrids such as the autumnal moth (Epirrita autumnata autumnata) from Europe or the hemlock looper (Lambdina fiscellaria fiscellaria) from North America, but rather a gradient outbreak insect.     

Oviposition by autumn gum moth (Mnesampela privata) in relation to Eucalyptus globulus defoliation, larval performance and natural enemies

Abstract 1 Eucalyptus globulus Labill. exhibited consistent intraspecific variation in oviposition choice by Mnesampela privata (Guenée) (Lepidoptera: Geometridae) in field surveys of host plants that had been designated, based on the prior season's defoliation levels, as resistant or susceptible to M. privata. At both field sites significantly fewer egg batches were found on resistant compared to susceptible trees. 2 In cage bioassays we demonstrated a significant oviposition preference by M. privata, with over two‐fold greater number of egg batches laid on foliage sprigs from susceptible compared to resistant trees. Despite differences in tree oviposition choice, we found no evidence in the field or the laboratory of adult females adjusting egg batch size in accordance to oviposition preference. 3 Caged larval survival, development times and pupal weight did not vary significantly between resistant and susceptible trees. Similarly, in a laboratory feeding experiment, neonates utilized resistant and susceptible foliage equally. 4 Larval mortality in the field attributed to natural enemies did not vary significantly between resistant and susceptible trees, nor did the percentage of Telenomus sp. parasitism of M. privata eggs within a batch and batches per tree. 5 Failure to associate either larval performance or natural enemy efficacy with the observed intraspecific variation in E. globulus susceptibility to M. privata oviposition indicates that some other unidentified factors drive the evolution of host selection for oviposition.     

Intra-plant host selection, oviposition preference and larval survival of Chrysophtharta agricola (Chapuis) (Coleoptera: Chrysomelidae: Paropsini) between foliage types of a heterophyllous host

Abstract 1 Paropsine chrysomelid beetles defoliate commercial eucalypt plantations in Australia. Adults and larvae feed on the same host, with the larval food source determined by the oviposition choice of females. Most eucalypt species are heterophyllous, with their foliage undergoing distinct morphological and chemical changes between adult and juvenile growth. 2 The intra‐plant foliage feeding and oviposition preference adults and the larval development of Chrysophtharta agricola were examined using adult and juvenile foliage of a heterophyllous plantation species, Eucalyptus nitens. The foliage types differ in chemistry, toughness, waxiness and timing of production. 3 In the field, feeding damage caused by adult beetles was 15% more frequent on adult foliage than on juvenile foliage; however, egg batches were three times more common on juvenile than on adult foliage. 4 Oviposition preference for juvenile foliage over adult foliage was confirmed in choice trials in the laboratory, with adult fecundity and longevity not significantly different between foliage types. 5 Larval survival, development time and subsequent pupal weight were also unaffected by foliage type, suggesting that neither foliage type is nutritionally superior for adults or for larvae. However, adult foliage was significantly thicker than juvenile foliage and this may prove a physical constraint to larval establishment. Biotic and abiotic factors (including interactions with natural enemies, competition, microclimate and mate location) that may affect patterns of host plant utilization are discussed.     

The influence of leaf age on the oviposition preference of Chrysophtharta bimaculata (Olivier) and the establishment of neonates

1 The degree of discrimination shown by a herbivore when selecting oviposition sites has been suggested as a key factor to understanding herbivore population dynamics. Chrysophtharta bimaculata (Coleoptera: Chrysomelidae) is a primary pest of Tasmanian eucalypt forests and can cause severe defoliation. Previous work suggests that females show discrimination when selecting oviposition sites. Our aim was to test the degree of oviposition discrimination exhibited by C. bimaculata with regards to leaf toughness, a character that is critical to neonate survival. 2 We conducted an experiment examining the leaf toughness critical for neonate survival and found that significant larval mortality occurs above a toughness of 46.9 g. We also determined that the maximum toughness of leaves upon which larvae established in the field was 48.2 g, supporting the laboratory result. 3 Field surveys showed that although the majority of eggs were laid on leaves suitable for larval establishment, many eggs were laid on unsuitable, tougher leaves. However, all eggs were normally placed within 20 cm of suitable leaves and glasshouse trials demonstrated the neonates could move this distance without mortality occurring. 4 We conclude that egg batch distribution and larval performance of C. bimaculata will influence the population dynamics of C. bimaculata in two ways. Firstly, the availability of expanding/newly expanding leaves of eucalypt hosts will determine larval carrying capacity. Secondly, at a more localized level, the deposition of large numbers of egg batches on both suitable and unsuitable leaves followed by successful neonate migration increases the risk of resource depletion and poor larval development.     

Outbreak of an invasive paropsine beetle in south‐west Ireland: preference,performance and damage to Eucalyptus

An exotic, paropsine beetle –Paropsisterna nr. gloriosa Blackburn – occurred at high densities in south‐west Ireland in 2007. In bioassays, adults and larvae fed on foliage from a variety of eucalypt species. Eggs and neonates occurred only in association with new foliage. Despite their ability to consume old foliage, adult beetles had a high preference for new leaves with low specific leaf weights (softer leaves). In choice tests, adults that depleted new foliage of their preferred host, moved to new foliage of a second host but not older foliage of the preferred host. A 2008 survey of southern Ireland indicated that P. nr. gloriosa was restricted to County Kerry, largely associated with foliage plantations in that county. The distribution of damage suggests that the initial spread of the beetle was facilitated by foliage‐trade activities. Eucalyptus parvula L.A.S. Johnson & K.D. Hill was the most heavily damaged species at many plantations. Eucalyptus pulverulenta Sims and Eucalyptus cordata Labill. were highly resistant to the beetle as indicated by low levels of damage in the field and reduced fitness of larvae in feeding trials. Nevertheless, at the plantation with the highest overall levels of damage, adult beetles moved to feed on E. pulverulenta. A progressive dispersal from plantations also caused slight damage to neighbouring ornamental eucalypts. Clear preferences by P. nr. gloriosa for new foliage, irrespective of eucalypt species, suggests that pollarding – the removal of top branches to produce dense juvenile foliage – accelerated population build‐up during 2007.     

Variation in the ability of larvae of phytophagous insects to develop on evolutionarily unfamiliar plants: a study with gypsy moth Lymantria dispar and Eucalyptus

• 1 By examining variation in the abilities of polyphagous insects to develop on host plants with secondary metabolites that they have never encountered previously, we may be able to gain some insights into the nature of evolution of biochemical mechanisms to process plant secondary metabolites by phytophagous insects.
• 2 The present study aimed to examine variation in the ability of gypsy moth larvae Lymantria dispar (Lymantriidae) to complete development on different species of the plant genus Eucalyptus (Myrtaceae). Leaves of at least some Eucalyptus species contain formylated phloroglucinol derivatives. These are secondary metabolites that are evolutionarily unfamiliar to the gypsy moth.
• 3 Larvae of gypsy moth showed extremely variable responses in larval performance between Eucalyptus species, between individual trees within host plant species, between moth populations, and between individuals within moth populations.
• 4 Larval survivorship was in the range 0–94%, depending on the host. Failure of at least some larvae to complete development on some Eucalyptus species indicates that gypsy moth larvae have a limited ability to process secondary metabolites in eucalypt leaves.
• 5 At least some individuals, however, appear to already possess biochemical mechanisms that process the secondary metabolites in leaves of Eucalyptus species, and therefore the abilities of larvae to complete development on phylogenetically and chemically unfamiliar hosts are already present before the gypsy moth encounters these potential hosts.

     

Larval performance in relation to oviposition site preference in olive kernel moth (Prays oleae Bern., Yponomeutidae,Praydina)

1 The tri‐voltine moth Prays oleae Bern. spends its larval stages on the native olive tree (Olea europaea L. var. sylvestris Brot. and five cultivars, Oleaceae) mining the leaves, the flowers and the fruits in each generation; it seldom switches to other native or introduced confamilial plant species. 2 In this study the pattern of oviposition of the olive moth was examined in olive fields and natural vegetation, in relation to in situ recruitment as an outcome of processes such as density dependence or risk spreading. 3 Larval body size (width of epicranial sclerites) was also examined and compared between host substrates, years and morphological, physiological, ecological and nutritional attributes of the host. 4 The factors influencing the oviposition pattern of the olive moth such as the carbon/nitrogen ratio, number of flowers, branch length and previous leaf damage were ranked differently in different cultivars. 5 ‘Hot spots’, i.e. olive trees or parts of trees receiving a high egg load, were found to be the result of in situ recruitment. 6 Physiological mortality among first instar larvae was significantly negatively correlated with the number of oviposited upon leaves; suggesting that the adult selects for oviposition the best available substrate. 7 As adult moths selected leaves with minimal probability of abscission for oviposition, leaf abscission was not a major mortality factor, although first instar larvae reduced leaf longevity. 8 Host quality did not affect all larval stages in the same way. 9 The more nutritionally poor the substrate, the longer the duration of the larval stage feeding on it. The phenological timing of the insect life stages very closely tracks the phenological phases of its host plant, primarily focusing on the most nutritious host stage in terms of larval performance.     

Variable responses of tuber moth to the leaf trichomes of wild potatoes

This study examines the response of tuber moth, Phthorimaea operculella (Zeller) (Lepidoptera: Gelechiidae), during the initial stages of attack, to variability in trichome density and composition on foliage of Solanum berthaultii (Hawkes) and Solanum tarijense (Hawkes) (Solanaceae). Solanum berthaultii bears two types of glandular trichome (type A and type B) that together reduced oviposition by the moth. Females were often completely deterred from ovipositing on foliage with >300 trichomes per cm². In contrast, neonate establishment on S. berthaultii was generally positively related to trichome densities, indicating that trichomes may be a poor defense against P. operculella when the moth oviposits in soil and neonate larvae select the host plant. Solanum tarijense has only one type of glandular trichome (type A) and eglandular hairs. Most eggs were deposited on the adaxial leaf surfaces that had lower trichome densities. Although the density of type A trichomes was negatively related to oviposition, high densities of hairs on the abaxial and adaxial leaf surfaces appeared to stimulate oviposition, leading to stronger positive relations between hair densities and oviposition. Larvae generally established on the abaxial surface where hair densities were greatest. Relationships between the abaxial densities of leaf hairs and neonate establishment on S. tarijense were positive. The results indicate that the responses by P. operculella to the types and density of trichomes are complex. Whereas type A and type B trichomes may act synergistically to reduce oviposition by the moth, leaf hairs do not defend against oviposition and neither leaf hairs nor type A and B trichomes reduce neonate establishment by this herbivore species.     

Rapid wound-induced resistance in white birch (Betula pubescens) foliage to the geometrid Epirrita autumnata: a comparison of trees and moths within and outside the outbreak range of the moth

Two strains of a geometrid defoliator, Epirrita autumnata, were used in bioassays to test existence and relative efficacy of rapid, wound-induced foliage resistance in two provenances of the white birch. One birch and one moth strain originated in the outbreak range of the moth and another outside it. Both birch provenances responded to manual leaf damage by changes in foliage quality which significantly retarded growth of the insects, reducing their pupal weights and protracting larval periods. Leaves which were previously damaged were lower quality as Epirrita food than adjacent intact leaves. Both of them were lower quality than intact leaves without damaged leaves nearby. Because of variance between years in the efficacy of the response, and because of different transfer distances of the provenances to the common garden where the experiments were performed, we could not ascertain whether there is any overall difference in the efficacy of rapid inducible responses between the provenances. Both moth strains were affected by wound-induced deterioration in foliage quality. There were no differences in how the moth strains experienced inducible resistance in the two birch provenances. Moths achieved relatively higher pupal weights on the birch provenance matching their origin. Moths from the outbreak range completed their larval period in a shorter time and pupated in a smaller size and, due to dependence of fecundity on size, had a lower potential rate of increase than insects outside the outbreak range.     

Impact of reassociation with a coevolved herbivore on oviposition deterrence in a hostplant

Although selection by herbivores for increased feeding deterrence in hostplants is well documented, selection for increased oviposition deterrence is rarely examined. We investigated chemical mediation of oviposition by the parsnip webworm (Depressaria pastinacella) on its principal hostplant Pastinaca sativa to determine whether ovipositing adults choose hostplants based on larval suitability and whether hostplants experience selection for increased oviposition deterrence. Webworms consume floral tissues and florivory selects for increased feeding deterrents; moths, however, oviposit on leaves of pre-bolting plants. Exclusive use of different plant parts for oviposition and larval feeding suggests oviposition should select for increased foliar deterrents. Recent webworm colonization of New Zealand (NZ) allowed us to assess phenotypic changes in foliar chemicals in response to webworm oviposition. In a common garden experiment, we compared NZ populations with and without a history of infestation from 2004 to 2006 for changes in leaf chemistry in response to oviposition. Three leaf volatiles, cis- and trans-ocimene, and β-farnesene, elicit strong responses in female moth antennae; these compounds were negatively associated with oviposition and are likely oviposition deterrents. Leaf β-farnesene was positively correlated with floral furanocoumarins that deter florivory; greater oviposition on plants with low floral furanocoumarins indicates that moths preferentially oviposit on parsnips most suitable for larval growth. Unlike florivory, high oviposition on leaves did not lower plant fitness, consistent with the fact that NZ parsnip foliar chemistry was unaffected by 3–6 years of webworm infestation. Thus, in this system, selection by ovipositing moths on foliar chemistry is weaker than selection by larvae on floral chemistry.     

Glandular trichomes of Solanum berthaultii and its hybrids with potato deter oviposition and impair growth of potato tuber moth

Glandular trichomes on foliage of the wild potato species, Solanum berthaultii Hawkes, deter oviposition by the potato tuber moth (PTM), Phthorimaea operculella Zeller and negatively affect other important performance parameters. Oviposition deterring factors are localized in the glandular trichomes of S. berthaultii. When mechanically transferred to foliage of a susceptible potato cultivar, trichome contents reduced egg laying by 97%. Removal of glandular trichomes from S. berthaultii foliage using a combination of chemical and mechanical procedures increased oviposition rates ca. 210-fold. Removal of trichomes also led to increased mobility of larvae on the leaf surface, more leaf feeding, shorter larval development and larger pupae. The resistance conferred by glandular trichomes of S. berthaultii provides an important genetic trait potentially useful for management of PTM.     

Leaf age and life history variables of a leafminer: the case of Liriomyza trifolii on potato leaves

Field observations and laboratory experiments have demonstrated that infestation by the serpentine leaf miner, Liriomyza trifolii Burgess (Diptera: Agromyzidae), begins in the lower leaves of the potato plant, and proceeds through the middle leaves to those of the upper canopy. In choice and no‐choice experiments, mated adult L. trifolii females were given access for 48 h to potato plants, and caged on differently aged leaves. The extent of their feeding and oviposition on the 5–7 leaflets of the upper, middle, and lower leaves were recorded. The life history variables of the next generation were estimated: percentage egg hatch, number of mines formed, larval survival, number of pupae formed, size and weight of pupae, percentage pupation, number of adults formed, percentage emergence, size and weight of adults, sex ratio, adult longevity, and their reproductive performance. The results showed that L. trifolii females laid fewer eggs on the upper leaves, which were poor hosts for larvae. However, a comparison of oviposition behavior between the middle and lower leaves showed that the data did not fit the oviposition preference–offspring performance hypothesis, which postulates that females preferentially oviposit on hosts on which larvae perform best. Females exhibited a preference for the larger, older, lower leaves, although the middle leaves were superior for the growth and development of the young stages. It is hypothesized that adult ovipositional preference for the older, larger, and thicker leaves of the lower foliage may be influenced by factors other than resource quality for larvae.     

Host selection,oviposition behaviour and leaf traits in a specialist willow sawfly on species of Salix (Salicaceae)

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Biology of the Eucalyptus leaf beetle Paropsisterna selmani (Chrysomelidae: Paropsini): a new pest of Eucalyptus species (Myrtaceae) in Ireland

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Reduction of oviposition time and enhanced larval feeding: two potential benefits of aggregative oviposition for the viburnum leaf beetle

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Oviposition preference of a Eucalyptus herbivore and the importance of leaf age on interspecific host choice

1. Paropsine chrysomelids are eucalypt folivores for which it has been hypothesized that the availability of suitable age class foliage (i.e. young foliage) is a more important determinant of host use than the physical and/or chemical characteristics of foliage from different hosts. This prediction was tested by assessing the oviposition preference of Chrysophtharta bimaculata (Olivier) on excised branches of Eucalyptus regnans F. Mueller and E. nitens (Deane & Maiden) Maiden.
2. In cage trials, and as observed in the wild, beetles oviposited preferentially on E. regnans when the branches of both eucalypt species displayed the full range of foliar developmental classes, from immature to fully expanded. However, when the branches were pruned so that both species displayed only immature and expanding leaves, no difference in oviposition between the species occurred. Further, when the branches were pruned so that E. regnans displayed only fully expanded leaves and E. nitens displayed only immature and expanding leaves, females oviposited entirely on the latter species.
3. These results support and extend the hypothesis that C. bimaculata 's ovipositional preference is dictated more by leaf age class than by the tree species from which the leaves derive. In the discussion it is proposed that a eucalypt that initiates and expands leaves rapidly will be less preferred for oviposition than one that develops and expands leaves slowly, because of the smaller area of its canopy with less sclerophyllous foliage.     

Seasonal allocation of defense investment in Ilex opaca Aiton and constraints on a specialist leafminer

Summary Phytomyza ilicicola (Diptera: Agromyzidae), a univoltine specialist leafminer, is one of the few insect herbivores of American holly. Adult emergence is closely synchronized with leaf flush in spring, and females make numerous feeding punctures on and oviposit in new leaves. Larvae hatch in late May and June, but their feeding period and development are prolonged so that more than 80% of the mine enlargement occurs from January until March of the following year. We propose that this unusual life cycle reflects adaptation to constraints imposed by seasonal and age-related changes in chemical and structural defenses, and in nutritional quality of holly foliage. As holly leaves age, there is a shift in allocation of defense investment away from allelochemicals, including phenolic compounds and saponins, toward leaf sclerophylly, spinose teeth, and low foliar nitrogen and water. Rapid increases in leaf toughness and decreases in nutritional quality limit availability of leaf tissues for adult feeding and oviposition to a two-to threeweek phenological window during leaf flush. Mature holly foliage is a nutritionally poor resource by nearly all criteria known to affect food quality for herbivores. This may be the main reason for the prolonged larval development of P. ilicicola. Alternatively, winter feeding and pupation in spring may be adaptations which help to ensure synchrony of adult emergence with leaf flush. Repeated puncturing by female P. ilicicola does not render leaves more suitable for larvae, nor is it a means by which females sample leaf exudate to assess leaf quality prior to oviposition. Rather, leaf puncturing occurs mostly on leaves that are relatively high in soluble nitrogen, and is apparently a means by which females obtain protein and sugars prior to and during oviposition.The investigation reported in this paper (No. 85-7-8-208) is in connection with a project of the Kentucky Agricultural Experiment Station and is published with the approval of the Director     

Suitability of Eucalyptus and Corymbia for Mnesampela privata (Guenée) (Lepidoptera: Geometridae) larvae

Abstract 1 Mnesampela privata (Guenée) has a host list of 40 Eucalyptus and at least one Corymbia species. Larval survival and performance was studied on 19 species to investigate how certain leaf traits influence the suitability of different species. 2 After 7 days, survival on Eucalyptus aggregata and Eucalyptus camphora is greater than 70% even though the toughness of leaves is 0.15–0.19 mg/mm². However, after the same time, survival on genotypes of Eucalyptus melliodora and Eucalyptus sideroxylon was less than 60%, or even 0%, even though the toughness of some leaves was as low as 0.11 mg/mm². An unmeasured allelochemical, rather than toughness, may reduce survival on these species. 3 Dry weights of first‐instar larvae were negatively correlated with leaf toughness for 13 of the species studied. Species that produced the heaviest first‐instar larvae were not the same hosts that produced the heaviest second‐instar larvae. 4 Dry weights of female pupae were negatively correlated with total oil content for five of the species studied. 5 Larvae exhibit age‐related changes in feeding behaviour. Neonates skeletonize leaves (avoid leaf veins and oil glands) and post‐third‐instar larvae ingest whole leaf fragments (consume small leaf veins and oil glands). These findings suggest that neonates are sensitive to high leaf toughness and non‐oil plant secondary metabolites whereas older larvae are less sensitive to high leaf toughness and are likely to become larger adults on hosts with lower oil contents.     

The chemical world of crucivores: lures, treats and traps

Papilio polytes utilizes only a few plant species of Rutaceae as hosts in the field. We examined in detail the acceptability of Toddalia asiatica (a major host plant) and three other potential rutaceous hosts, Murraya paniculata, Melicope triphylla, and Phellodendron amurense, for ovipositing females of the butterfly. Female responses to the foliage, methanol extracts, and partitioned fractions from these plants were assayed for the presence of oviposition stimulants and/or deterrents. Larval survivorship on these plant species was also compared as an estimate of fitness. The foliage and a methanol extract of T. asiatica readily induced egg-laying, while females responded moderately to the foliage and a methanol extract of P. amurense. By contrast, ovipositing females only marginally accepted Me. triphylla and completely rejected Mu. paniculata. Further experiments to test the biological activity of fractions derived from the respective plant species revealed that T. asiatica contains potent oviposition stimulant(s) and that weak stimulant(s) are present in P. amurense. Poor or negative oviposition responses to both Me. triphylla and Mu. paniculata proved to be attributable to strong deterrent(s) present in these plants. Larvae performed very well on T. asiatica and P. amurense, whereas larval mortality was much higher on Mu. paniculata and Me. triphylla, suggesting the involvement of antifeedant(s) or toxic substance(s) in these plants.     

Flexible patch time allocation by the leafminer parasitoid, Opius dimidiatus

Abstract.

• 1 The ability to use flexible decision rules can be an advantage to parasitoid females searching for patchily-distributed hosts. In a series of laboratory experiments the hypothesis that Opius dimidiatus, a solitary parasitoid of the chrysanthemum leafminer (Liriomyza trifolii), adjusts the time she allocates to searching for her larval hosts in response to both patch qualities and experiences with hosts was tested by varying such patch parameters as area, presence of host mines and density of host mines, and by allowing ovipositions and encounters with parasitized hosts.
• 2 Though leaf area was not a factor, the presence of host mines in a leaf did increase the time a female O.dimidiatus spent searching, over time spent on unmined leaves.
• 3 When host mine density was increased, females responded by increasing their search period in a density-dependent manner, suggesting a perception of patch quality.
• 4 Ovipositions in hosts caused females to reset their‘giving-up time’(GUT), or increase search intensity, by adding an amount of search time that increased with each successive oviposition. Conversely, encounters with parasitized (unsuitable) hosts incremented the GUT, but by an amount that decreased with each successive encounter.