Colonization, survival, and causes of mortality of Camerariu hamadryadella (Lepidoptera: Gracillariidae) on four species of host plants

Abstract.

• 1 The abundance, survival, and causes of mortality of Cameraria hamadryadella (Clemens) (Lepidoptera: Gracillariidae) were examined on four host plant species in Virginia, U.S.A. Quercus alba L. and Q.rubra L. are native within the geographic range of C.hamadryadella, and Q.robur L. and Q.benderi Baenitz are exotic. Q.robur is native to Europe, North Africa, and Asia and was probably introduced prior to 1850, and Q.benderi is of hybrid origin and introduced to cultivation before 1900. Q.alba and Q.robur are in the subgenus Lepidobalanus (white oaks), and Q.rubra and Q.benderi are in the subgenus Erythrobalanus (red oaks).
• 2 Larval mines of C. hamadryadella were abundant on both white oak species, including the exotic Q.robur, but were rare on host plants in the red oak subgenus. Un-hatched eggs of C.hamadryadella were not observed on red oaks. Other observations on host distribution indicate that C.hamadryadella is rarely found on red oaks. These observations are interpreted as circumstantial evidence that C. hamadryadella generally avoids ovipositing on red oaks.
• 3 Survival of C.hamadryadella to the adult stage was similar among all host species, but larvae tended to survive longer on hosts in the red oak subgenus. The observation of higher survival rates of early instar larvae on red oaks suggests that no nutritional or secondary chemical barrier reinforces the observed pattern of oviposition.
• 4 Significant differences in the distribution of the causes of mortality were detected between native and exotic host plant species. Larvae and pupae on native hosts were more likely to die because of predation, while those on exotic host plants were more likely to die because of parasitism and host feeding by adult female parasitoids. This pattern could arise because parasitoids prefer to forage on exotic host plants or because predators avoid foraging on exotic plants.
• 5 This study shows for C. hamadryadella that the only barriers to colonization and use of exotic hosts, in the white and red oak subgenera, are the presence of cues sufficient to stimulate oviposition and/or the absence of cues to deter oviposition. It also suggests that the presence of closely related native host plants in the region of introduction will increase the probability that exotic plants will be colonized by phytophagous insects.

     

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.

     

Aggregation in field parasitoid populations: foraging time allocation by a population of Diadegma (Hymenoptera, Ichneumonidae)

ABSTRACT.

• 1 A field study was made of foraging time allocation by a population of parasitic wasps, Diadegma spp. (Ichneumonidae), to plants containing different densities of their hosts, the caterpillars of Plutella xylostella (L.).
• 2 The parasitoid population exhibited a clear aggregative response, spending more total time on higher density patches, which probably resulted from wasps making more and longer visits to these densities.
• 3 Despite this aggregation, positive density dependent parasitism was not found. The functional response of the Diadegma population exhibited an upper asymptote at high host densities, probably due to an increase in the proportion of time spent handling hosts, which countered the effect of aggregation.
• 4 While Diadegma may select and forage preferentially on plants with higher host density, they do not exhibit the tendency, predicted by some optional foraging models, to exploit progressively less profitable plants during a foraging bout. Some factors affecting patterns of parasitoid foraging in the field are discussed.

     

The effects of host plant phenology on the demography and population dynamics of the leaf-mining moth, Cameraria hamadryadella (Lepidoptera: Gracillariidae)

Abstract.

• 1 We examined the effects of variation in the timing of spring leaf production and autumn leaf fall on the survival, mortality and abundance of Cameraria hamadryadella on Quercus alba and Q.macrocarpa.
• 2 We monitored and manipulated the timing of foliation on field and potted Q.alba trees and observed the abundance of C.hamadryadella on those trees. We also monitored and manipulated the timing of leaf fall on Q.alba and Q.macrocarpa trees in the field and observed its effects on survival, mortality and abundance of C.hamadryadella.
• 3 Variation in the timing of spring leaf production has no effect on C. hamudryadella abundance. However, a warm winter and spring in 1991 led to accelerated development and the imposition of a facultative third generation in one out of ten years of observation.
• 4 In 1989, leaves fell relatively early and leaf fall in the autumn accounted for more than 50% of the mortality of C.hamudryadella. in 1990 and 1991 leaves fell relatively late and leaf fall induced mortality was substantially reduced and overwinter survival was markedly increased.
• 5 The abundance of C.hamadryadella remained constant in the spring and summer of 1990 following the previous autumn's relatively early leaf fall, but increased by 10-fold in the spring of 1991 following the relatively late leaf fall of autumn 1990. The abundance of C.hamadryadella also increased 4-fold between the summer of 1991 and the spring of 1992 after another autumn of relatively late leaf fall. We attribute these increases in abundance in part to reduced mortality because of later leaf fall.
• 6 Variation in the timing of autumn leaf fall may be responsible for initiating outbreaks of C.hamadryadella.

     

Host-attacking behavior of a eulophid parasite,Kratochviliana sp., to the leaf mining host,Phytomyza ranunculi (Diptera: Agromyzidae) during its stay on the leaf

The present paper studies how the female parasite of Kratochviliana sp. visits and attacks its host larvae of Ranunculus leaf mining fly, P. ranunculi at a single leaf visit. The parasite visited its hosts at random on the leaf. The frequency of host visits was independent of the host density and the proportion of hosts survived from the parasite attack, in a leaf and its distribution was expressed as a single straight line. It almost always attacked living hosts at the first host visit after isolated from them for one day but with the rate of about 0.5 at the subsequent visits. In consequence, the relationships of the number of host attacks and killed hosts to the host density drew satulated curves in each. A model of host attack by this parasite at its single leaf visit was formulated by modifyingBakker et al.'s model (1972) basing upon these observations and the attack avoidance by the parasite to already attacked hosts previously reported.     

Defence and development in a gregarious leaf-mining beetle

Abstract.

• 1 The gregarious larvae of the chrysomelid beetle Microrhopala vittata mine the leaves of goldenrods (Solidago spp.). These mines serve both as food and as shelter for the larvae.
• 2 Life-table data and experiments indicated that mine initiation and moves to secondary mines represented especially vulnerable stages during larval development. Leaf mines effectively protected M.vittata against predators in the field.
• 3 Field experiments indicated that larvae hatching from larger clutches of eggs stood a greater chance of surviving to pupation, primarily because larvae hatching in groups proved more successful at initiating leaf mines. Once inside the leaf mine, however, larvae feeding in large groups attained lower adult masses, and were more likely to abandon the natal mine and did so earlier in development because large groups more rapidly destroyed a leaf.

     

Density-dependent regulation of Aedes cantans (Diptera: Culicidae) in natural and artificial populations

Abstract.

• 1 Mortality and size variation (siphon length) in Aedes cantans larvae were examined in natural populations in northern England in 1989 and 1990.
• 2 Under crowded conditions, density-dependent competition led to reduction in the size of both larvae and adults and increased larval mortality.
• 3 Larvae were also maintained in cages in the field at different densities. Results paralleled those for the natural populations in the ponds; larvae maintained at high densities showed increased mortality and reduced size.
• 4 Possible density-dependent factors leading to mortality and size reduction include cannibalism and contact inhibition leading to food shortages.
• 5 The main density-independent factor contributing to larval mortality was habitat desiccation.

     

Host location by the fruit fly parasitoid Diachasmimorpha krausii: role of fruit fly species,life stage and host plant

• 1 Diachasmimorpha krausii is a braconid parasitoid of larval tephritid fruit flies, which feed cryptically within host fruit. At the ovipositor probing stage, the wasp cannot discriminate between hosts that are physiologically suitable or unsuitable for offspring development and must use other cues to locate suitable hosts.
• 2 To identify the cues used by the parasitoid to find suitable hosts, we offered, to free flying wasps, different combinations of three fruit fly species (Bactrocera tryoni, Bactrocera cacuminata, Bactrocera cucumis), different life stages of those flies (adults and larvae) and different host plants (Solanum lycopersicon, Solanum mauritianum, Cucurbita pepo). In the laboratory, the wasp will readily oviposit into larvae of all three flies but successfully develops only in B. tryoni. Bactrocera tryoni commonly infests S. lycopersicon (tomato), rarely S. mauritianum (wild tobacco) but never C. pepo (zucchini). The latter two plant species are common hosts for B. cacuminata and B. cucumis, respectively.
• 3 The parasitoid showed little or no response to uninfested plants of any of the test species. The presence of adult B. tryoni, however, increased parasitoid residency time on uninfested tomato.
• 4 When the three fruit types were all infested with larvae, parasitoid response was strongest to tomato, regardless of whether the larvae were physiologically suitable or unsuitable for offspring development. By contrast, zucchini was rarely visited by the wasp, even when infested with B. tryoni larvae.
• 5 Wild tobacco was infrequently visited when infested with B. cacuminata larvae but was more frequently visited, with greater parasitoid residency time and probing, when adult flies (either B. cacuminata or B. tryoni) were also present.
• 6 We conclude that herbivore‐induced, nonspecific host fruit wound volatiles were the major cue used by foraging D. krausii. Although positive orientation to infested host plants is well known from previous studies on opiine braconids, the failure of the wasp to orientate to some plants even when infested with physiologically suitable larvae, and the secondary role played by adult fruit flies in wasp host searching, are newly‐identified mechanisms that may aid parasitoid host location in environments where both physiologically suitable and unsuitable hosts occur.

     

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.

     

Searching at different spatial scales: the foraging behaviour of the aphid parasitoid Aphidius rosae in rose bushes

The searching patterns of the aphid parasitoid Aphidius rosae were analysed at three different spatial scales: leaves, shoots and bushes. Parasitoid females searched aphid-infested leaves and shoots intensively and remained on average more than twice as long on infested than on uninfested shoots. Patch times and oviposition numbers per shoot were highly variable both between females and for different shoot visits within females. However, at the shoot and bush level low oviposition numbers were generally found. The time spent on different behavioural patterns (searching, resting, feeding, host handling) changed significantly during subsequent shoot visits of individual females but oviposition success was not influenced by this change. Parasitoids searched individual leaves and shoots mainly by walking, while moving between shoots occurred exclusively by flight. The travel time between shoots (i.e. flight time) accounted for less than 1% of the residence time in a bush. At the bush level foraging was characterized by a high ability to localize infested shoots and consequently little time was wasted in searching on uninfested shoots. The pattern of resource exploitation of individual females was consistent with the distribution of A. rosae larvae in field samples taken from individual rose bushes.     

Time allocation of the parasitoidAphidius colemani (Hymenoptera: Aphidiidae) foraging forAphis gossypii (Homoptera: Aphidae) on cucumber leaves

The time allocation of individualAphidius colemani female parasitoids foraging forAphis gossypii nymphs on cucumber leaves has been investigated. Apart from experiences on the current leaf (such as density of hosts on the current leaf, density of hosts on a neighboring leaf and encounters with hosts on the current leaf), the effect of previous leaf visits on the time allocation was studied. Behavioral records were analyzed by means of the proportional hazards model, to determine the tendency of leaving the current leaf. The leaving tendency decreased only on leaves with a high host density (100 aphids), thus increasing the giving up time since the latest encounter. Rejection of aphids had no influence on the leaving tendency. To assess the effect of the number of hosts encountered on the leaving tendency, we considered three classes: 0–30 encounters, 31–100 encounters, and 100 or more encounters with hosts. The effect of the number of hosts encountered differed at different aphid densities. When fewer than 10 aphids were present the leaving tendency was much greater after 30 encounters than beforehand. At a density of 100 aphids the leaving tendency was lower than at the other aphid densities and increased only after 100 encounters. The density of hosts on a neighboring leaf, ranging from 0 to 100 hosts, had a negligible effect on the leaving tendency. Repeated visits to leaves with 10 unparasitized aphids resulted in an increase in the leaving tendency after 10 visits. It is argued that the parasitoids have some innate expectancy of host availability and that they concentrate on high-density patches.     

Population dynamics of the yew gall midge Taxomyia taxi and its chalcid parasitoids: a 24-year study

Abstract.

• 1 The dynamics of three populations of Taxomyia taxi (Inchbald) (Diptera: Cecidomyiidae) and its chalcid parasitoids have been studied over a 24-year period. Most individuals have a 2-year life cycle but some develop in 1 year. Details of within-generation mortalities in T.taxi are used for life table analyses.
• 2 Mortality in the period between emergence of adult T.taxi and larval infestation of buds appears to be density-dependent and is the largest component of overall mortality. In 2-year life cycles, this mortality and that caused by Torymus nigritarsus (Walker) contribute equally to variance in overall mortality. In 1-year cycles, mortality caused by Mesopolobus diffinis (Walker) is density-dependent and accounts for most within-gall losses.
• 3 T. nigritarsus, which attacks only 2-year galls, is absent from all populations for a number of years in the middle of the study period. Its searching efficiency declines as its density and that of its host increase.
• 4 Densities of M.diffinis are strongly correlated with those of available hosts. Percentage parasitism of 2-year galls is less than that of 1-year galls, suggesting the occurrence of enemy-free space.
• 5 Although there are some correlations in densities and mortality between trees, the dynamics of populations on each are frequently different.
• 6 An earlier analysis of 10 years' data failed to demonstrate density-dependent effects. The extension to a 24-year run has revealed such effects, but also demonstrates long-term fluctuations in population densities, with corresponding changes in the balance of advantage between 1- and 2-year life cycles of T.taxi.

     

Host population structure affects field sex ratios of the heteronomous hyperparasitoid, Coccophagus atratus

Abstract.

• 1 Second instar Filippia gemina de Lotto scale insects are the preferred hosts of female Coccophugus atrutus Compere larvae. These scale insects were found on their host plants, Chrysanthemoides monilifera Norlindh and Cliffortia strobilifera Mettenius, only at certain times during a 1 year sampling programme.
• 2 Late larval instars and prepupae of C.atratus, and a Metaphycus species, are the preferred hosts of male C.atratus larvae. These hosts, although they occurred on the same host plants as hosts for female C.atratus, were most numerous at different times during the sampling period.
• 3 The ratio of hosts suitable for C.atratus varied from a predominance of hosts suitable for females through to a predominance of hosts suitable for males. Sex ratios of adult C.atratus followed a similar trend but did not reflect, exactly, the ratio of available hosts. Differences in mortality between sexes and hyperparasitism may account for this anomaly.
• 4 Variable population sex ratios observed in C.atratus apparently result from the behaviour of individual females in which brood sex ratios are dependent on the relative availability of hosts for males and hosts for females. This behaviour, in turn, may result from variability in the host population structure but may also result from selection pressures operating at the time that heteronomous hyperparasitism evolved.

     

The effects of foliage damage on casebearing moth larvae, Coleophora serratella, feeding on birch

ABSTRACT.

• 1 Mechanical damage to birch (Betula pendula Roth) leaves leads to an increase in the concentration of phenolic compounds, which spreads throughout the leaf within 8 days.
• 2 Coleophora serratella L. (Lepidoptera: Coleophoridae) apparently responds to this chemical change over a similar time scale. Within 24 h of pin-pricking leaves the casebearer moves from the immediate vicinity of the damage, but is just as likely to move to an undamaged portion of the damaged leaf as to vacate the leaf entirely. After 8 days mines on undamaged portions of damaged leaves were significantly smaller than mines on undamaged leaves.
• 3 Furthermore, Coleophora serratella reared on damaged trees took an average of 3 days longer to develop than those reared on undamaged trees.
• 4 It has been suggested that increased movement in response to damage-induced chemical changes causes hyperdispersed damage on plant foliage. Both within and between-leaf casebearer damage patterns were shown to be aggregated on birch.
• 5 Thus although mechanical damage can induce chemical and behavioural changes in the field, these are not reflected in the observed damaged patterns. We speculate on several possible reasons for this.

     

Targeted sugar provision promotes parasitism of the cereal leaf beetle Oulema melanopus

• 1 Parasitoids may often lack access to sugar (e.g. floral nectar) in agricultural settings. Strategically timed spraying of host plants with sugar solution may provide one means of enhancing parasitism at the same time as minimizing nontarget effects (e.g. benefiting the pest itself).
• 2 Sucrose was sprayed in wheat fields of northern Utah (U.S.A.) to assess the effects on parasitism of the cereal leaf beetle Oulema melanopus by the larval parasitoid Tetrastichus julis.
• 3 Early‐season sugar provisioning, when larvae of the pest were first hatching and parasitoid adults were newly emerged, did not affect the numbers of cereal leaf beetle larvae that matured in treated plots but increased parasitism rates of beetle larvae by four‐fold in 2006 and by seven‐fold in 2007.
• 4 No net influx of adult parasitoids into plots was detected after the application of sugar. Locally‐emerging parasitoids may have spent less time searching for their own food needs versus hosts. A laboratory experiment also confirmed that access to sucrose significantly increased parasitoid longevity.
• 5 The field experimental results obtained demonstrate that applications of sugar, implemented to target a key time of the growing season when benefits are maximized for parasitoids and minimized for their hosts, can strongly promote parasitism of the cereal leaf beetle in wheat fields.

     

Non-random distribution patterns of leaf miners on oak trees

Summary Leaf-mining Stilbosis quadricustatella larvae are distributed non-randomly within leaves of their host plants, sand live oak (Quercus geminata) and water oak (Q. nigra), in north Florida. Fewer mines are found together on the same side of the mid-vein than separated, on opposite sides of the mid-vein. Larvae do not normally cross the mid-vein but create small blotch-like mines along subsidiary veins. Investigations of the usual mortality factors acting on these leaf-miner populations, including competition, parasitism, and predation, revealed no significant differences in these factors between mines separated by the mid-vein and those on the same side of the leaf. However, early leaf abscission, which kills the larvae present in the leaf, occurs significantly more frequently in cases where larvae are clustered on one leaf side. The reasons for this differential leaf abscission are not yet clear.     

Recruitment of native parasitoids by an exotic leaf miner, Cameraria ohridella: host-parasitoid synchronization and influence of the environment

Abstract

• 1 The horse‐chestnut leaf miner, Cameraria ohridella, is a moth of unknown origin that has recently invaded Europe and severely defoliates the European horse‐chestnut, an important ornamental tree.
• 2 Several indigenous parasitoids have colonized this new host, but parasitism remains low. One of the hypotheses suggested to explain the low parasitism is that candidate parasitoids emerge too early in spring to attack the first host generation and, thus, need early‐occurring leaf miners as alternate hosts. This hypothesis was tested by observing the synchronization between the phenology of the moth and that of its main parasitoids, and by comparing parasitism rates and parasitoid richness in different environments with various levels of biological diversity.
• 3 In spring, the bulk of the parasitoids emerge at least 5 weeks before the occurrence of the first suitable larvae of C. ohridella whereas most parasitoid adults reared outdoors die within 5 weeks after emergence.
• 4 Parasitism rates and parasitoid richness do not increase with biological diversity, suggesting that most parasitoids attacking the first generation of C. ohridella do not come from alternate hosts. Parasitism does not increase later in the year in the subsequent generations, when host‐parasitoid synchronization becomes less critical.
• 5 We conclude that, although the spring emergence of parasitoids is not synchronized with the phenology of C. ohridella, the parasitoids attacking the first generation are probably old or late‐emerging adults of the overwintering generation. The lack of synchronization is probably not the only reason for the poor recruitment of native parasitoids by C. ohridella.

     

Host-age discrimination during host location byCotesia glomerata, a larval parasitoid ofPieris brassicae

Some parasitoids are restricted with respect to the host stage that they attack and even to a certain age within a stage. In this paper we investigate whether the parasitoidCotesia glomerata can discriminate between old and young caterpillar instars of its host,Pieris brassicae, before contacting these hosts, since contacts with older instars are very risky with a chance of being killed, due to the aggressive defensive behaviour of the caterpillars. Flight chamber dual choice tests showed that volatile chemicals emitted by Brussels sprouts plants (Brassica oleracea var. gemmifera) after feeding damage by 1st and 5th larval instars are equally attractive to the wasps. Simulated herbivore damage by 2nd and 5th larval instars, obtained by treating mechanically damaged leaves with carterpillar regurgitant, was also equally attractive, even when the wasps were exposed to repeated experience on different larval instars to increase their discriminatory ability. In contrast, single choice contact bioassays showed that the time spent searching on a leaf with feeding damage of 1st instar larvae was significantly longer than the time spent on 5th instar feeding damage or on mechanically damaged leaves. Both flight and contact bioassays did not show any effect of egg-related infochemicals. The results demonstrate thatC. glomerata can discriminate between young and old larval instars ofP. brassicae, without contacting the caterpillars. This is not done through volatile herbivore-induced synomones but through cues that are contacted after arrival at a caterpillar-infested leaf.     

Proximate effects of maternal oviposition preferences on defence efficacy and larval survival in a diet‐specialised tortoise beetle. Who knows best: mothers or their progeny?

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Oviposition patterns and larval damage by the invasive horse‐chestnut leaf miner Cameraria ohridella on different species of Aesculus

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