Foraging behavior of a Dipteran leaf miner on exploited and unexploited hosts

Summary The foraging behavior of females of the leaf miner, Agromyza frontella (Rondani), (Diptera: Agromyzidae) when encountering unexploited or exploited alfalfa plants was studied in large field cages and in laboratory bioassays. Females did not recognize any exploited leaflets before contacting them and did not distinguish between leaflets with an egg or first instar larva and unexploited leaflets, even after contact. Only one fly oviposited in leaflets which contained 80–120 nutrition holes, one late second or third instar larva or which were marked with an epideictic pheromone in field cages. In laboratory bioassays females oviposited less in leaflets containing a second or third instar larva or an empty larval mine than in unexploited ones. Females foraging on unexploited leaflets engaged in area-restricted search and 10 of 11 females remained on the test plant for the full 60 min of observation. However, females foraging on exploited plants were much more active, spent a greater proportion of their time searching for suitable hosts, had the highest rates of visitation to all above ground plant parts and emigrated to the cage walls before 60 min had elapsed. These quantitative measures of foraging behavior indicated that females ranked plants after landing on them in the following order: unexploited plants >plants marked with pheromone or with many nutrition holes >plants with late instar larvae. The order of host ranking by foragers was in general agreement with the suitability of the host plants for larval survival, development and reproduction, as estimated from previous laboratory studies.Females of A. frontella foraging on unexploited alfalfa plants fed and oviposited significantly more often in the upper apical leaflets than in the lower, older leaflets. However, the choice of feeding site by flies on exploited plants did not vary with leaflet position (age), indicating that females fed in order to sample leaflet quality and that females investigated lower (older) leaves after they discovered that the preferred upper leaves were occupied. These data suggest that high quality oviposition sites may be limiting for A. frontella females, which could explain why superparasitism of leaflets sometimes occurs in nature, even when unexploited sites are available.     

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.

     

Larval parasitoids and pathogens of the groundnut leaf miner,Aproaerema modicella (Lep.: Gelechiidae), in India

Natural enemies of the groundnut leaf miner,Aproaerema modicella (Deventer), were studied at the International Crops Research Institute for the Semi-Arid Tropics (ICRISAT) located near Hyderabad in peninsular India. Hymenopterous parasitoids attacking leaf miner larvae were the most important group of natural enemies. Nine primary and eight secondary parasitoids emerged from host larvae, and killed up to 50% of the leaf miner larvae sampled. The trophic relationships between primary and secondary parasitoids are incompletely understood. The influence of pathogens of this species is reported for the first time. These pathogens killed up to 30% of the leaf miner larvae. The combined effects of all mortality agents killed up to 95% of the leaf miner larvae per sample period. However, use of insecticides in sprayed plots reduced the efficacy of parasitoids. The impact of predators on larval populations was not studied and may explain underestimates of leaf miner mortality rates.      

Changes in spatial distribution pattern during the larval stage of the fall webworm,Hyphantria cunea drury (Lepidoptera: Arctiidae)

Summary The changes in spatial distribution pattern during larval stage of the fall webworm,Hyphantria cunea were quantitatively investigated in the field experimental populations. The female adult deposits eggs as a cluster and the hatchlings make a compact colonial-web. In this period, the all-or-none type mortality which is characteristic in gregarious insect species was occasionary recognized before spinning a compact colonial-web. Once making a compact colonial-web, the larvae feed the leaves in the colonial-web up to about 5th instar. In this period, the movement of larvae occurred due to the local food shortage in a colonial-web and the expansion of colonial-web. As the larvae developed, the colonial-web was separated into several small groups. These larvae began to disperse about 5th instar. In this period, the local food shortage seems to be an important trigger for the larval dispersal. The mean concentration of larvae on leaves abruptly decreased, and finally the larvae became solitary at the 6th or 7th instars. The dispersal process in later larval stage is not necessarily due to the complete food shortage. The dispersal prior to the occurrence of food shortage may be a safety mechanism to protect the larvae from the food shortage.     

Willow leaf traits affecting host use by the leaf-gall-forming sawfly

Gall-forming insects usually have very restricted host ranges, but plant traits affecting patterns of host use have rarely been examined. The sawfly Phyllocolpa sp. (Hymenoptera: Tenthredinidae) forms leaf-roll galls on three of seven Salix (Salicaceae) species that occur together on riverbanks in central Japan. We have attempted to explain this host-use pattern by invoking three plant traits: synchrony of leaf flush with the oviposition period of the sawfly, intrinsic leaf quality as a potential larval food, and leaf morphology. Two Salix species frequently used by the sawfly, Salix eriocarpa and Salix pierotii, had similar leaf traits suitable for larval survival. The third species, Salix serissaefolia, was used relatively less often and the sawfly frequently stopped laying eggs on the plant during oviposition, suggesting ovipositional selection. S. serissaefolia had the smallest leaves, and survival of sawfly larvae was lower on S. serissaefolia than on S. eriocarpa and S. pierotii, because of gall destruction, by other herbivorous insects, and leaf-size restrictions. Among the four unused species, Salix chaenomeloides had a late leaf-flush phenology, Salix gracilistyla had inferior leaf quality, and Salix gilgiana had linear leaves; these seemed to be critical factors for non-use. Salix subfragilis was also unused, but the reason for this could not be explained by the three leaf traits studied.     

On interference among larvae of the greasy cutworm,Agrotis ypsilon rott. (lepidoptera: Noctuidae)

Summary The greasy cut worm,Agrotis ypsilon Rott., has a characteristic habit; the larva feeds on the small seedling of the host plant. Studies in a population in outbreak area suggested that the food shortage temporally took place, causing the reduction of weights of pupae and adults. The fact that this species depends on the small seedling as the host seemed to be responsible for this food shortage. However, it was found that the larval crowding also affected the body weight, even if the food was supplied in excess. It was further suggested that the weight reduction is due to the injury caused by the encounter of larvae. The oviposition habit to lay small egg masses and the aggressive behaviour of the larva was considered to be responsible for the uniform distribution at the later larval stage. Probably, the larval dispersion enabled the larva to forage in the most effective way as well as it protected the larva from the attack of another ones. As a result, this would enable the maximum number of individuals to survive. It seemed that the functions noted above did not work in the population being under outbreak condition, because the larval density was unusually high due to mass-invasion of moths.     

The change of the distribution pattern of the large weevil,Hyposipalus gigas Fabricius (Coleoptera,Rhynchophoridae) within a single generation. A preliminary note

The distribution pattern of adults of the large weevil, Hyposipalus gigas on logs of the Japanese black pine and that of larvae in pine stumps were examined by using the m^*−m method (Iwao , 1968). The distribution of adults on logs was contagious, probably due to their response to local heterogeneity of the environment. The larval distribution was also contagious, but there was no significant correlation between the number of larval bores and the stump size. The mortality of young larvae seemed not to be related with the number of larvae per stump, but it was related with the density per unit area of bark surface. There was some sign of inversely density-dependent or ‘all-or-none’ type mortaltiy in the late larval stage.     

A study of natural populations of the celery leaf-miner,Philophylla heraclei L. (Diptera,Tephritidae) II. Importance of changes of mines for larval populations

The leaf-mining larvae of the celery fly, P. heraclei, have the ability to leave their primary mine to bore a secondary mine in another leaflet or leaf. This phenomenon is always numerically significant under natural conditions (depending on the plots and the years, 38 to 97% of the larvae observed made such a change), although it is not an obligatory behavior. The distribution of the secondary mines on the leaflets and on the leaves varies with the importance of the migrations, with the total unoccupied and healthy leaf area and with the relative position of different leaves on the plant. These migrations are due to the insufficiency of healthy food available for the larvae: either because the quantity of parenchyma offered by a leaflet is too small for a larva, or because there is intraspecific competition following multiple egg-layings on the same leaflet, or because there is a deterioration of the quality of the parenchyma, particularly following the development of celery leaf-spot. The possibility for the larvae of P. heraclei to bore secondary mines appears to be an extremely important factor in the population dynamics of this species. This permits the reduction of the negative effects of intraspecific competition and celery leaf-spot, and permits the colonization by the larvae of young leaves not used by the females at the time of egg-laying.     

Oviposition,larval survival and leaf damage by the willow leaf blotch miner,Micrurapteryx salicifoliella,in relation to leaf trichomes across 10 Salix species

1. Foliar trichomes clearly reduce chewing damage and efficiency of movement by some insect herbivores, but the effect of trichomes on insect oviposition is less well characterised. Trichomes are likely to have particularly strong, negative effects on species that require secure attachment of the egg to the leaf epidermis for successful transition to the feeding stage – a group that includes many leaf mining insects. 2. One such species, Micrurapteryx salicifoliella, must initially enter leaf cells directly from an egg adhered to the cuticle, but later instars can move between leaves and initiate new mines from the leaf exterior. 3. Natural patterns of occurrence by M. salicifoliella were quantified on 10 sympatric Salix species varying in trichome expression to test whether trichomes were associated with reduced oviposition, larval survival and leaf damage. 4. Mean egg density and leaf mining damage were negatively related to mean trichome density across Salix species. Survival of M. salicifoliella from egg to pupa was positively related to trichome density, suggesting that initiation of new mines by late‐instar larvae was not adversely affected by trichomes. There was no evidence that trichomes benefited leaf miner larvae indirectly by decreasing density‐dependent mortality; rather, the positive relationship between trichome density and larval survival may reflect less effective chemical defence by Salix species expressing high trichome density. 5. The results suggest that foliar trichomes serve as an effective defence against M. salicifoliella by deterring oviposition, but do not reduce the survivorship of those individuals that successfully transition from egg to larva.     

Dynamic interaction between a leaf beetle, Galerucella nipponensis, and an aquatic plant, Trapa japonica. II. Dispersal behavior of larvae

The dispersal behavior of the larvae of a leaf beetle, Galerucella nipponensis, was investigated to analyze the dynamic interaction between the leaf beetle and its aquatic host plant, Trapa japonica. The hatchlings dispersed from the hatched leaf to other leaves in a rosette within 12 h. Some larvae even dispersed on the water surface. As a result, the hatchlings distributed uniformly within the rosette. This larval dispersal appeared to be adaptive behavior responding to a rapid turnover of T. japonica leaves. The female adults preferred young or middle-aged leaves to old leaves as oviposition sites. Received: June 14, 2001 / Accepted: December 17, 2001     

Vascular implications of Dasineura sp. galls' establishment on Peumus boldus stems

• Some chewing larvae are capable of inducing galls in the host vascular cylinder, e.g. Dasineura sp. (Cecidomyiidae) on Peumus boldus stems. Due to the medicinal and economic importance of P. boldus, the anatomical and functional implications of establishment of Dasineura sp. on P. boldus stems were investigated. We asked if establishment of Dasineura sp. in P. boldus stems induces abnormalities at the cellular and organizational level of the vascular system that increase during gall development in favour of the hydric status of the gall.
• Anatomical alterations induced in the stems during gall development were determined. Cytohistometric analyses in mature galls were compared to non-galled stems, and water potential and leaf area of non-galled stems were compared with galled stems.
• Dasineura sp. establishes in the vascular cambium, leading to delignification and rupture of xylem cells, inhibiting formation of phloem and perivascular sclerenchyma. Gall diameter increases together with larval feeding activity, producing a large larval chamber and numerous layers of nutritive tissue, vascular parenchyma, and sclerenchyma. These anatomical alterations do not affect the leaf area of galled stems but favour increased water flow towards these stems.
• The anatomical alterations induced by Dasineura sp. in P. boldus stems guarantee water and nutrient supply to the gall and larva. After the inducer exits stems, some host branches no longer have vascular connections with the plant body.

     

Population regulation of a mosquitoArmigeres theobaldi with description of the animal fauna in zingiberaceous inflorescences

1. Inflorescences of some Curcuma and Zingiber (Zingiberaceae) in tropical Asia provide an unique aquatic habitat being discrete, small, and made of numerous smaller compartments (the bracts).
2. The aquatic community in inflorescences of Curcuma in northern Thailand was composed of immature Diptera, of which the biting midge Dasyhelea and the mosquito Armigeres theobaldi were the commonest. No important competitors, predators or pathogenic parasites for the mosquito were confirmed.
3. Inter-inflorescence distribution of the mosquito was contagious. Within each inflorescence, the fourth-instar larvae or pupae usually occupied bracts singly.
4. The k-value analysis detected density-dependent mortality due to contest competition in the mosquito larvae.
5. Variations in the larval and pupal mosquito size were density-independent and remarkably small as compared with size variations known for other mosquitoes.
6. These population attributes (large density-dependent mortality with density-independent, minimally variable individual size) appear unique among mosquitoes, arising from conspecific killing, efficient foraging (inter-bract movement by crawl and single occupation of bracts), and availability of host plant tissues as supplementary food.
7. A simple population model suggested that a small proportion of adult females lay eggs.

     

Leaf miner-induced changes in leaf transmittance cause variations in insect respiration rates

Very little is known about alterations in microclimate when an herbivore feeds on host plant. Modifications of leaf transmittance properties induced by feeding activity of the leaf miner Phyllonorycter blancardella F. were measured using a spectrometer. Their effects on the herbivore's body temperature and respiration rate have been determined under controlled conditions and varying radiation level employing an infrared gas analyser. By feeding within leaf tissues, a miner induces the formation of feeding windows which transmit a large portion of incoming radiations within a mine. As a result, body temperature and respiration rate increase with radiation level when positioned below feeding windows. Therefore, the miner is not always protected from radiations despite living within plant tissues. The amount of CO(2) released by larvae below feeding windows at high radiation levels is about five-fold that recorded in the dark. By contrast, body temperature and respiration rate increase only slightly with radiation level when the insect is positioned below intact tissues through which radiation is only weakly transmitted. A mine offers its inhabitant a heterogeneous light environment that allows the insect larva to thermoregulate through behavioural modification. Our results highlight the importance of physical feedbacks induced by herbivory which alter significantly an insect's metabolism independently of its nutritional state.     

Larval cannibalism during the late developmental stages of a facultatively gregarious encyrtid endoparasitoid

• 1 The larvae of many gregarious parasitoid species are usually non‐aggressive when they develop in or on a host, but those of Metaphycus flavus are one of the few exceptions known. Herein we describe their aggressive behaviour and the conditions under which it occurs, using observations in which larval development and physical conflict within parasitised and superparasitised hosts were mapped daily.
• 2 Metaphycus flavus larvae often engaged in physical conflict that resulted in consumption of the losing larvae (= cannibalism ) in superparasitised hosts, whereas such conflict and consumption occurred rarely when a single brood developed in a host.
• 3 Cannibalism among M. flavus larvae only occurred after the host resources had become scarce. Typically it occurred after the sixth day of development (fourth‐instar larvae) when the larvae in a clutch had separated from their aeroscopic plate and were freed of their attachment to the host's cuticle.
• 4 Female larvae in the initial clutch appeared more aggressive than male larvae when a second clutch was allocated 4 h after the first clutch. The probability of a larva being attacked and consumed by a brood mate increased as the number of larvae increased in the host. This partial tolerance might allow the members of the initial brood to defend themselves from offspring of a superparasitising female (= competitors ). Such post‐ovipositional regulation of brood size might be interpreted as high‐density intolerance among female offspring.

     

Clutch size in a leaf-mining fly (Pegomya nigritarsis: Anthomyiidae)

ABSTRACT.

• 1 Pegomya nigritarsis Zetterstedt is a gregarious leaf-mining fly attacking Rumex sp. The larvae from one clutch of eggs all inhabit the same mine.
• 2 Egg-batch size varies between host plants and is correlated with leaf size.
• 3 On Rumex obtusifolius L. larval survival as a function of clutch size was measured in the field. This measure was incorporated in an optimal clutch size model and a clutch size prediction obtained.
• 4 The predicted optimal clutch size, three, is that most frequently found in the field.

     

Leaf age and larval performance of the leaf beetle Paropsis atomaria

ABSTRACT.

• 1 Larval performance of the leaf beetle Paropsis atomaria Oliver was determined for larvae raised on both new and mature leaves of Eucalyptus blakelyi Maiden. Larvae were transferred to mature leaves at different ages; control larvae stayed on new leaves through all instars.
• 2 Only larvae reared on new leaves through the third instar survived to pupate on mature leaves; developmental time was prolonged by 20% and pupal weight was reduced by 50% in these larvae compared with larvae reared entirely on new leaves. Almost all larvae died when transferred to mature leaves as first, second or third instars.
• 3 Low survival and slow development on mature leaves was mainly due to failure by larvae to feed. Larvae palpated leaves and could discriminate among leaf ages immediately, without biting into the leaf tissue.
• 4 New leaves had higher concentrations of oil and tannins than old leaves, while there were no significant differences in nitrogen concentrations in the two types of leaves. Mature leaves were more than 3 times tougher than new leaves, in terms of g mm^?2 of penetrometer force.
• 5 In drought years E. blakelyi may not produce sufficient new leaves to supply specialist herbivores with their preferred food resource. We infer that drought years reduce P. atomaria larval performance significantly, and influence the population dynamics of the insect.

     

Host plant, oviposition behavior and larval ecology of a sawfly leafminer, Profenusa japonica (Hymenoptera: Tenthredinidae)

The host plant, oviposition behavior and larval ecology of Profenusa japonica Togashi are reported for the first time. Adults of P. japonica mated and oviposited on a polyantha rose, Rosa multiflora (Rosaceae), in April. Each female adult laid an egg on the edge of a leaflet. Hatched larvae consumed the parenchymatous layer of leaflets and in so doing created a mine. The larval stage comprised five instars. On average, 70.6% of the total area of a leaflet was consumed by one larva. Female adults of P. japonica laid eggs singly, probably to avoid larval competition for food.     

Variation in rates of leaf abscission between plants may affect the distribution patterns of sessile insects

Summary At Sand Lake, Leon County, Florida, mines of Stilbosis quadricustatella, a leaf-mining moth, occur on sand live oak trees (Quercus geminata) over a broad range of densities. Some trees have fewer than 2% of their leaves mined (lightly infested), others up to 70% (heavily infested). Similar levels of infestation are maintained on the same trees year after year. There are no significant negative correlations of miner density per tree with any mortality factor that might explain miner preferences for certain trees. Nor is there a positive correlation with host leaf quality as measured by foliar nitrogen or by total or individual amino acid concentrations of host leaves. Egg-transfer experiments showed that larvae from eggs transferred to lightly infested trees were more likely to be killed by leaf abscission than were those that were transferred to and developed on heavily infested trees. This is the first demonstration that variation in rates of leaf abscission could be an important cause of the observed distribution pattern of sessile insects between conspecific host plants.     

The distribution of risk and density-dependent mortality in the galls of Eurosta solidaginis, the goldenrod gall fly

ABSTRACT.

• 1 This paper explores the net effect of a suite of mortality factors on a sedentary prey, the larvae of the goldenrod gall fly, Eurosta solidaginis Fitch (Diptera: Tephritidae).
• 2 Mortality is caused by unknown factors early in larval development, two species of parasitoid wasp (Hymenoptera: Eurytomidae), an inquiline beetle larva (Coleoptera: Mordellidae), and during the winter months downy woodpeckers Picoides pubescens (L.).
• 3 Distribution of mortality among galls relative to prey (gall) distribution was measured and discussed with respect to the distribution of relative risk of predation.
• 4 Galls are by and large contagiously distributed among quadrats, and mortality is distributed in a comparable pattern to that of galls.
• 5 The pattern of mortality on Eurosta larvae is neither density-dependent nor aggregated independently of gall distribution. Persistence in the system is probably a result of a combination of other factors such as adult mortality and early larval death which may have intergenerational density-dependent effects, and the linkage of locally unstable sub-populations via migration.

     

The population dynamics of Cheilosia fasciata (Diptera,Syrphidae): significance of environmental factors and behavioural adaptations in a phytophagous insect

Summary A population of Cheilosia fasciata Schin. & Egg., 1853 was studied in a beech forest (Melico-Fagetum allietosum) near Göttingen (FRG). This syrphid species is closely associated with ramson (Allium ursinum L.), as the larvae mine the plant's leaves. Adult abundance was determined using emergence traps (1981–1986); egg and larval abundance was determined by examining ramson leaves: puparia were extracted from soil samples (1984 and 1986). Cheilosia fasciata is a univoltine species; the imagines emerge in late April/early May. Larval development took five (1986) to six (1984) weeks. In late June/early July the larvae enter the soil for pupariation. A linear relationship was found between the area of the mines and larval biomass (dry wt.). It was concluded that one larva had to mine 51.90 cm² of leaf area in order to gain the mean maximum dry weight (11.15 mg). Furthermore, it was shown that ovipositing females, laying just one egg per leaf, tended to select large leaves providing this minimum amount of food. It is hypothesized that females probably follow straight-lined routes on their oviposition flights, rather than ovipositing on leaves chosen at random. By applying such a strategy, females can almost completely avoid competition for food among their own offspring. Furthermore, competition among the progeny of the whole population is also reduced, particularly in years when adult abundance is very low. The significance of certain factors influencing the population dynamics of Cheilosia fasciata was evaluated by applying a rank correlation analysis. It was shown that high precipitation rates in April reduced the time available for mating and oviposition resulting in low emergence abundance in the next spring. Larval survivorship was enhanced by high precipitation rates in May and June, indicating that drought may be unfavourable to larvae and reduce food availability. Predation by a parasitoid Phygadeuon ursini Horstmann, 1986 (Hymenoptera, Ichneumonidae), appears to be just an imperfect (Milne 1984) density-dependent control mechanism; in fact, it was shown that food limitation is the only significant mechanism of density-dependent population control.