Leaf trenching by Indian red admiral caterpillars for feeding and shelter construction

Many folivorous insects sever leaf veins, trench across leaf blades, or girdle petioles prior to feeding on leaves. The purpose of these behaviours is generally thought to be sabotage of the anti-herbivore chemical defences of host plants. For insects that construct leaf shelters, these behaviours may make leaves easier to roll or fold. Larvae of the Indian red admiral butterfly, Vanessa indica (Herbst), cut trenches at the base of host plant leaves and construct leaf-fold shelters. I attempted to determine which of the above functions the trenching behaviour of V. indica larvae serves. If the function of trenching is to facilitate leaf-folding, then larvae should cut trenches before folding leaves. When the larvae used larger leaves, they tended to trench them prior to folding them. This suggests that the function of trenching behaviour is to make leaves easier to fold. However, larvae sometimes cut trenches after folding the leaves, suggesting that this behaviour has additional functions. When the larvae were about to moult or pupate, they did not cut trenches, suggesting that trenching behaviour may also be involved in feeding.     

Fitness costs and benefits of shelter building and leaf trenching behaviour in a pyralid caterpillar

1. Shelter building and petiole trenching in the Lepidoptera is a behaviour that mediates ecological pressures including those exerted by both food plants and natural enemies. 2. Fitness costs and benefits of trenching and shelter‐building behaviour related to predation and larval performance were investigated in a pyralid species that inhabits and feeds on leaf shelters. 3. Assays comparing the performance of caterpillars feeding on trenched versus non‐trenched foliage and fresh versus dry leaves were conducted. Whereas pupal weight was positively affected by petiole trenching, larval development was delayed when caterpillars fed on dry leaves. 4. A field experiment comparing predation on caterpillars inside and outside shelters demonstrated that predation was significantly higher for exposed caterpillars. 5. No physiological costs associated with shelter building were found given that caterpillars performed equally regardless of the number of shelters they built. 6. The effect sizes of top‐down and bottom‐up forces on pupal weight, development time, and predation risk indicated that the major effect of shelters is through the reduction of predation risk. The integration of experiments and natural history observations showed that fitness benefits provided by shelters change across ontogeny.     

Predation and avoidance of tough leaves by aquatic larvae of the moth Parapoynx rugosalis (Lepidoptera: Pyralidae)

Abstract.

• 1 Aquatic larvae of the pyralid moth Parapoynx rugosalis Möschler repeatedly construct protective cases by cutting portions (discs) from leaves of the waterlily Nymphaea ampla and sandwich themselves between the disc and the underside of the host leaf. Construction of a new case requires leaving the old case, thus increasing exposure to predators and parasites.
• 2 In an experiment, larvae with protective cases experienced no mortality due to predation by fish, whereas larvae without cases experienced substantial predation.
• 3 In a series of choice tests, larvae preferentially selected young, tender leaves over old, tough leaves for construction of cases, and larvae spent significantly less time completing their shelters when cutting discs from young, tender leaves.
• 4 A partial explanation of why larvae select young, tender leaves for construction of their protective shelters may be that exposure time to predators during construction is minimized.
• 5 The same mechanism linking preferences for tender leaves to reduced exposure to predators during construction may also apply to other insect herbivores exhibiting leaf-rolling or case-building behaviour.

     

Roles for structural and temporal shelter-changing by fern-feeding lepidopteran larvae

Summary Larvae of the pyralid moth, Herpetogramma aeglealis, construct feeding shelters upon the Christmas fern (Polystichum acrostichoides). Field and laboratory study involving 532 shelters showed that as the larvae mature, they sequentially inhabit approximately 5 shelters of 3 distinct types, constructed at night on different fronds of the same plant. The bundle shelter, simple and ephemeral, is first to be inhabited and constructed. The fiddlehead shelter which houses slightly older larvae strongly resembles contemporaneously emerging frond fiddleheads. The final shelter form, the globe, is a silk-bound ball of leaflets at the frond tip. An individual larva usually constructs 3 globe shelters on different fronds of the same plant before completing its development. As shelter sites, sterile Polystichum fronds are chosen preferentially over fertile fronds. The bundle and fiddlehead shelter forms, less abundant, appear cryptic to humans and perhaps to other vertebrates. The final globe shelter form is larger and quite conspicuous. However, the persistence of empty globe shelters left on the plant as the larva moves to a new one may serve to make searching for larvae less profitable for potential predators and parasites. We suggest that the energetic costs of constructing and occupying multiple shelters may be offset by circumvention of reduced frond palatability and reduced exposure to predators and parasites.     

Ecological consequences of shelter sharing by leaf-tying caterpillars

A wide variety of insect herbivores construct and inhabit leaf shelters (ties, rolls, folds, and webs). Shelter construction can lead to a high rate of secondary occupation by other arthropods, including other species of constructors. The consequences for the inhabitants of secondarily occupying these shelters are currently unknown. In this study, we conducted field experiments to examine the fitness consequences (survival and attack by natural enemies) for caterpillars that (i) occupy a shelter with conspecifics vs. occur singly; and (ii) establish a new shelter vs. colonize a pre‐existing one. In addition, we conducted factorial laboratory experiments to test the hypothesis that caterpillars sharing shelters with conspecifics might have reduced construction costs (a potential benefit of shelter‐sharing or secondary occupation). Larvae of Psilocorsis quercicella Clemens (Lepidoptera: Oecophoridae) placed in white oak [Quercus alba L. (Fagaceae)] leaf ties alone or in groups of three had equal likelihood of survival from natural enemies. This same caterpillar species, however, had a higher disappearance rate when placed in pre‐existing leaf ties than when placed in newly formed ones, suggesting a potential cost of secondary colonization. A similar experiment with a closely related species [Psilocorsis cryptolechiella (Chambers)], however, failed to detect a cost of secondarily occupying shelters made on beech, Fagus grandifolia Ehrh. (Fagaceae). In the laboratory experiment, we found no evidence of shelter‐sharing benefits; rather larvae reared in shelters in groups of three had lower pupal mass (and thus lower potential fecundity) than larvae reared singly, suggesting a cost of shelter sharing. Moreover, groups of larvae forced to repeatedly construct new shelters tended to have reduced survival relative to the other treatment, suggesting that energetic constraints are more likely to reduce fitness when larvae cohabit shelters. Taken together, these results indicate that the common phenomenon of shelter sharing by leaf‐tying caterpillars has either neutral or negative effects for the occupants. The fact that these leaf‐tying caterpillars actually share shelters may simply reflect limited availability of oviposition sites.     

Female's preference for oviposition site and larval performance in the water-lily beetle,Galerucella nymphaeae (Coleoptera: Chrysomelidae)

Water-lily beetles prefer younger rather than older water-lily leaves as oviposition sites. By the time of hatching, however, young leaves have aged consieerably. Larval performance of the water-lily beetle was measured on different types of leaves of the yellow water-lily and compared with oviposition preference of females. The leaf types used in the experiments were categorized as (i) young, (ii) natal (medium-aged) and (iii) old. The natal leaves were the ones on which larvae from a particular egg-batch had hatched. There were two sets of experiments. First, larvae were raised from eggs to pupae on young and on old leaves. Second, the growth of the 1st-instar larvae was measured on young, natal, and old leaves. The development time from egg to pupa did not differ between young and old leaves, but larvae growing on young leaves attained a higher pupal weight. In the second experiment the 1st-instar larvae grew fastest on their natal leaves, but there was also variation in the growth rate of progeny from different egg-batches. Larval growth on young and old leaves did not differ significantly. Larvae tried to emigrate much less from natal than other types of leaves. Females tended to lay eggs on leaves where larval growth was fastest. It seems that medium-aged leaves are best for larval growth, but the leaf characteristics responsible for this remain unresolved.     

Specialized generalists: constraints on host range in some plusiine caterpillars

Insects that feed on plants with secretory canals often cut trenches across leaves, thereby depressurizing the canals and eliminating exudation at their distal feeding site. We compared the trenching ability of three species of polyphagous plusiines (Lepidoptera: Noctuidae) and tested if trenching scores correlate with larval growth. The three plusiines (Trichoplusia ni, Pseudoplusia includens, and Rachiplusia ou) were each tested on prickly lettuce, Lactuca serriola (Asteraceae), which has latex canals, and on Italian parsley, Petroselinum crispum (Apiaceae) with oil ducts. To ascertain how secretory canals affect performance, larvae were tested on intact leaves and on excised leaves with depleted canals. T. ni larvae cut trenches in both plant species, whereas P. includens only trenched prickly lettuce and R. ou only trenched Italian parsley. Intact leaves of Italian parsley were acceptable to all three species. Trenching varied in T. ni and R. ou, but did not correlate significantly with larval growth. In contrast, trenching was required for feeding on intact prickly lettuce. Final-instar T. ni all cut trenches and developed rapidly. P. includens varied in trenching and performance; their trenching scores correlated with growth. R. ou larvae did not trench or feed even though most R. ou on intact Italian parsley cut at least partial trenches. All three plusiine species developed rapidly on excised leaves of both plant species, documenting the suitability of these plants when canals are inactivated. Our results document the efficacy of latex canals as a plant defense and suggest that trenching ability alone does not permit feeding. Larvae must also recognize the need to trench and must tolerate deterrent exudates during the trenching procedure. Received: 25 August 1999 / Accepted: 8 December 1999     

Uniformity of Leaf Shelter Construction by Larvae of Epargyreus clarus (Hesperiidae), the Silver-Spotted Skipper

Larvae of the silver-spotted skipper, Epargyreus clarus (Hesperiidae), construct shelters from leaves of their leguminous host plants, making four distinct shelter types that change predictably over larval ontogeny. Shelters built by first-instar larvae are located on the apical half of the leaflet and are almost invariant in size, shape, and orientation, suggesting a stereotypical process of shelter location and construction. We have determined that the regularity of these shelters results from a prescribed pattern of larval movements and behaviors, in which larvae use their body length as a ruler and employ silk not only as a building material but also as a template to guide the location of cuts in the leaf. Though lepidopteran larvae lack the sensitive antennae, long jointed appendages, and other measurement devices used by structure-building bees, wasps, and caddis flies, they can nonetheless use simple tools and behavioral patterns to produce characteristic and regular shelters.     

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.     

Preference for protected feeding sites by larvae of the willow-feeding leaf beetle Galerucella lineola

1. Free-living insects are often thought of as more vulnerable to environmental hazards than concealed insects, such as galling or mining insects. The possibility that larvae of the free-living leaf beetle Galerucella lineola seek out existing plant structures and thereby become partly concealed was explored.
2. Neonate larvae of G. lineola frequently feed in rolled-in margins of young leaves of their host plant, Salix viminalis . In addition to nutritional benefits from feeding on young leaves, larvae may also gain protection against adverse weather conditions and general predators by feeding in the leaf rolls. Field and laboratory experiments were conducted to test these hypotheses.
3. Artificial shelters were constructed and cohorts of neonate larvae were placed on experimental plants. In all experiments, larvae preferred to feed in shelters, even when shelters were constructed on mature leaves.
4. In one of the experiments, fewer larvae disappeared when shelters were provided. In a predator exclusion experiment, however, no differences in predator-inflicted mortality on G. lineola were found between shelter-containing shoots and control shoots.
5. A laboratory experiment showed increased protection from desiccation when shelters were present; growth rate was higher for larvae feeding on plants with shelters.
6. Thus, free-living insects may not always be as exposed to environmental hazards as is often assumed. In particular, young larvae may take advantage of preformed structures on their host plant and feed in a concealed microhabitat. Because mortality, in general, is high during early instars, shelter-seeking behaviour may increase survival significantly. The existence of preformed shelters may therefore be a plant characteristic that should be considered when exploring the environmental risks associated with the free-living habit.     

Ontogenetic timing of density dependence: location-specific patterns reflect distribution of a limiting resource

Theoretical considerations suggest that the relative abundance of age-specific limiting resources determines the ontogenetic timing of density dependence. Structural shelters may represent one such resource which can become increasingly scarce with increasing body size. Here we use a time series of juvenile Atlantic salmon (Salmo salar) densities and ask whether ontogenetic patterns of density-dependent losses in two separate reaches of a river can be predicted by considering their shelter abundances. The analyses were conducted using sampling site data (n = 30) as well as stream-reach averages. Loss rates from the egg to the young-of-the-year stage were density-dependent in both reaches. For the transition from the young-of-the-year to the yearling stage, when shelters are more likely to become limiting, the results were sensitive to the spatial scale of analysis. On the reach scale, among-year variation in loss rates was positively correlated with density in the reach with the lowest shelter abundance, whereas no such effect was found in the other reach. This demonstrates that the ontogenetic timing of density dependence can vary among areas within populations, and hence among populations, and that this variation can be explained by quantification of age-specific limiting factors. For analyses at the sample site scale this pattern was reversed, with stronger density dependence in the reach with highest shelter abundance. However, this result was clearly driven by immigration into low density sites, which masked the true reach-level effect. Thus, our study also exemplifies how population level regulation inferred from patch- or trap-based data that fails to account for animal movements can be biased.     

Effects of Leaf Age on Oviposition and on Offspring Fitness in the Imported Willow Leaf Beetle Plagiodera versicolora (Coleoptera: Chrysomelidae)

Imported willow leaf beetles Plagiodera versicolora oviposit on willow leaves, and both larvae and adults feed on the leaves. In the field, eggs were found on leaves near the center of branchlets, and the number of eggs per cluster was independent of the leaf area and position. However, in the laboratory, females chose young leaves over old leaves, for both oviposition and feeding and choice did not rely on information on relative position or size of leaves. Developing on young versus old leaves may provide both advantages and disadvantages. In the laboratory, larvae developed more quickly and attained a greater adult weight when fed young versus old leaves, perhaps because of increased mandibular wear of larvae fed old leaves. However, in the field, survival of eggs was lower on young versus old leaves. In the laboratory, rates of cannibalism and survivorship to adulthood did not differ on young versus old leaves.     

Contrasting engineering effects of leaf‐rolling caterpillars on a tropical mite community

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Spatial Orientation in Blattella germanica L. Larvae

The aim of these experiments was to investigate the type of cues used in homing processes by young Blattella germanica L. larvae. Several types of stimuli were tested: path integration with kinesthetic cues and visual orientation with landmark cues. Tests measured the escape direction of larvae from the food box after disturbance. Either type of cue alone, path integration or visual landmarks, was sufficient to allow larvae to orient towards their shelters, but they oriented more precisély when both types of cue were used. When several landmark cues (proximal and distal) were present, their relative angular position seemed important in the orientation process. Macroscopic shapes in the environment appeared to be used as a global image, memorized to reach the shelter.     

Sites chosen by diapausing or quiescent stage quino checkerspot butterfly, <Emphasis Type="Italic">Euphydryas editha quino</Emphasis>, (Lepidoptera: Nymphalidae) larvae

This study examines whether in nature endangered quino checkerspot (Euphydryas editha quino) larvae will return to diapause and if so where they choose to hide. Multiple years of diapause probably help larvae survive drought years and sites chosen have high survival value to the species. Ninety square meters of habitat were created by removing non native plants and replacing them with natives found at checkerspot occupied sites. During the 2005–2006 winter 1,000 post-diapause larvae were released. From these larvae 31 adults (20 males and 11 females) developed over a 2.5 month period (March 20–June 6) from 41 pupae. One chrysalis was parasitized by a parasitic wasp Pteromalus puparum (L.) in the family Pteromalidae, one was partially eaten by an animal, while the remaining eight pupae died of unknown causes. Thirty quadrats (1 square meter each) were cleared of vegetation, leaf and branch litter, rocks, and checkerspot larvae from July 5 to August 1, 2006. Forty-nine larvae were found that returned to diapause. Most larvae (31) chose to make shelters on California buckwheat, which is not a checkerspot food plant, two to five cm above the ground. One shelter had 22, another had seven, and two others had single larvae. Five of 10 larvae found in leaf litter below California buckwheat were crawling and not associated with shelters suggesting they had been dislodged from shelters. California buckwheat may be important in habitat restoration for the checkerspot, particularly at sites below 900 meters elevation where summer conditions are hot and dry. No additional larvae were found the following spring, when they should have exited diapause. Therefore 910 (91%) larvae were lost to some undocumented form of mortality.     

Movement and feeding patterns of Epilachna cucurbitae Richards (Coleoptera:Coccinellidae) on pumpkin and zucchini plants2

The oviposition patterns of adults and the movement and feeding patterns of larvae of Epilachna cucurbitae on two species of cucurbits, Cucurbita maxima cv Queensland Blue and C. pepo cv Blackjack, were studied in the field and laboratory. The physical and nutritional characteristics of host plant leaves of different ages were described. Younger leaves had higher nitrogen contents but were less abundant, smaller and had higher trichome densities than older leaves. The development of first instar larvae was delayed by the leaf hairs on young and mature pumpkin leaves which prevented larvae from reaching the leaf surface to feed First instal larvae developed more quickly on leaves rich in nitrogen. Neither the total developmental time of larvae nor the size of pupae was affected by leafage because larvae on poor quality leaves compensated by eating more. Female beetles oviposited on all but the youngest and oldest leaves of the host plant. The trichomes on young leaves prevented females from attaching eggs to the leaf surface. First instar larvae remained where they hatched, but older larvae were more mobile, Changing feeding sites frequently and moving progressively to younger, more nutritious leaves. Final instar larvae moved onto adjacent vegetation to pupate. The adaptive significance of these patterns is discussed in relation to the nutritional value, hairiness and abundance of host plant leaves of different ages and the physical limitations of different larval instars.     

Seasonal variation in leaf characteristics and food selectionby larval noctuids on an evergreen Mediterranean shrub

Despite year round availability of foliage, abundance of generalist noctuid larvae (Lepidoptera: Noctuidae) in evergreen-dominated Mediterranean forests has a narrow, distinct spring peak. This restricted larval period has been suggested to result in part from avoidance of the nutritionally poor mature foliage, and preference for nutritionally superior spring-produced young leaves. This study examines this hypothesis by (i) documenting differences in nutritional characteristics between expanding (April) and mature (June) young leaves of the evergreen Mediterranean shrub Daphne laureola L. (Thymelaeaceae), and (ii) experimentally studying the feeding preferences of noctuid larvae for young leaves, old leaves (≥ 1 yr old), and developing fruits of this species in one south-eastern Spanish locality. Young leaves of D. laureola declined in nutrient concentration and specific dry mass from April to June. The responses of noctuid larvae, in terms of both relative preference and total consumption, to this seasonal variation in chemical and physical features of young leaves were also investigated. When noctuid larvae were simultaneously offered young leaves, old leaves and developing fruits, they exhibited similar preferences for young leaves and developing fruits, and rejected old leaves developed during the previous year. Noctuid larvae did not modify their consumption of young leaves relative to old leaves and developing fruits in response to seasonal changes. Food selection patterns exhibited by D. laureola noctuid herbivores, notably the rejection of old leaves in favour of young ones, are consistent with the hypothesis relating restricted larval periods of these generalist consumers with the low food value of the previous season leaves of evergreen Mediterranean plants.     

Ontogenetic dietary changes of whitefish larvae: insights from field and experimental observations

Ontogenetic changes in resource use are widespread in many fish species. This study investigated the feeding habits of whitefish (C. lavaretus L.) larvae in Lake Annecy (France) coupled with experimental behavioral studies in order to identify the underlying mechanisms of the ontogenetic shifts in the diet. The predatory behavior of wild larvae, and the escape responses of their zooplankton prey were both videorecorded in experimental tanks under controlled laboratory conditions. Ontogenetic diet patterns showed that young whitefish larvae have a preference for small cyclops, while older larvae selectively predate cladocerans. Our experimental observations showed that the capture success rate also varied in relation to ontogenetic development in fish. Young larvae were more successful in capturing small copepods, whereas old larvae were more successful in capturing Daphnia. In addition, the larvae were able to adjust their predatory behavior (speed, pursuit) according to the swimming pattern of the prey. These observations suggest that the selective predation on cladocerans observed in old larvae is the outcome of both active and passive choices depending on the escape swimming behavior of the prey, and handling time of the predator.     

Plant exudates trigger leaf-trenching by cabbage loopers, Trichoplusia ni (Noctuidae)

Cabbage loopers, Trichoplusia ni, cut a narrow trench across leaves of plants that release exudate, then feed distal to the trench in an area of reduced exudation. The larvae do not normally trench plant species such as plantain, Plantago lanceolata, that lack exudate. To determine what cues elicit trenching, I reared larvae to the final instar on plantain, then applied test solutions to their mouthparts during feeding. Loopers that received latex from Lactuca serriola (Asteraceae) or phloem exudate from watermelon, Citrullus vulgaris (Cucurbitaceae), often responded by cutting a trench in plantain, even though these larvae had not previously encountered exudate nor previously trenched. Loopers that were allowed to trench and feed on L. serriola for 1 day prior to the assay subsequently cut trenches in plantain more frequently and in response to more fluids, including a viscous solution of polyethylene glycol and latex from a non-host, poinsettia (Euphorbia pulcherrima). Subsequent bioassays with larvae reared entirely on plantain tested whether bitter cucurbitacins or gelation are essential cues for trenching. Sap from non-bitter cucumber plants (Cucumis sativus) caused larvae to trench, showing that cucurbitacins are not required to induce trenching. Loopers also trenched after receiving cucumber sap that did not gel due to the addition of mercaptoethanol. An extract of sap lacking the proteins that cause gelation likewise triggered trenching. Further fractionation revealed that cucumber sap and also butternut squash sap (Cucurbita moschata) contain trenching stimulants that are small (molecular weight < 3,000) water-soluble molecules. Received: 13 January 1997 / Accepted: 6 June 1997