Host-plant effects on larval survival of a salicin-using leaf beetle Chrysomela aeneicollis Schaeffer (Coleoptera: Chrysomelidae)

Several species of willow leaf beetles use hostplant salicin to produce a defensive secretion that consists of salicylaldehyde. Generalist arthropod predators such as ants, ladybird beetles, and spiders are repelled by this secretion. The beetle larvae produce very little secretion when they feed on willows that lack salicylates, and salicin-using beetles prefer salicylate-rich willows over salicylate-poor ones. This preference may exist because the larvae are better defended against natural enemies on salicylate-rich willows. If this is true, the larvae should survive longer on those willows in nature. However, this prediction has not been tested. I determined the larval growth and survival of Chrysomela aeneicollis (Coleoptera: Chrysomelidae) on five willow species (Salix boothi, S. drummondiana, S. geyeriana, S. lutea, and S. orestera). These species differed in their salicylate chemistries and in leaf toughness but not in water content. The water content varied among the individual plants. Larval growth of C. aeneicollis did not differ among the five species in the laboratory, but it varied among the individual plants and it was related to the water content. In the field, C. aeneicollis larvae developed equally rapidly on the salicylate-poor S. lutea and on the salicylate-rich S. orestera. Larval survival was greater on S. orestera than on S. lutea in one year (1986), but there was no difference between them during three succeeding years. In another survivorship experiment, larval survival was low on the medium-salicylate S. geyeriana, but high on the salicylate-poor S. boothi and on S. orestera. Larval survival in the field was related to the larval growth and water content that had been previously measured in the laboratory. These results showed that the predicted relationship between the host plant chemistry and larval survival did not usually exist for C. aeneicollis. One possible reason for this was that the most important natural enemies were specialist predators that were unaffected by the host-derived defensive secretion. One specialist predator, Symmorphus cristatus (Hymenoptera: Eumenidae), probably caused much of the mortality observed in this study. I discuss the importance of other specialist predators to salicin-using leaf beetles.     

Plant wound compounds from oviposition scars used in host discrimination by a stem-galling sawfly

The stem-galling sawfly Euura lasiolepisuses one or more plant wound compounds resulting from oviposition scars as cues in host discrimination (avoiding sites occupied by conspecifics). Four experiments were conducted to test hypotheses about how Euurapartitions resources. Experiment 1 demonstrated that Euuraavoids ovipositing on nodes with scars from previous ovipositions. Experiment 2 showed no evidence that the sawfly uses oviposition-deterring pheromones and indicated there is a time lag following oviposition before the oviposition scar becomes a deterrent. Experiment 3 showed that sawflies avoid artificially formed scars, demonstrating that a plant cue alone can lead to host discrimination. Experiment 4 showed that visual or tactile cues are not necessary for host discrimination and indicated that a plant wound compound functions as an oviposition deterrent. Both experimental results and field surveys showed that Euuraoviposition scars were more uniformly distributed than expected if sawflies were ignoring previous ovipositions.     

Perfect is best: low leaf fluctuating asymmetry reduces herbivory by leaf miners

Fluctuating asymmetry (FA) represents small, random variation from symmetry and can be used as an indicator of plant susceptibility to herbivory. We investigated the effects of FA of two oak species, Quercus laevis and Q. geminata, and the responses of three herbivore guilds: leaf miners, gallers, and chewers. To examine differences in FA and herbivory between individuals, 40 leaves from each tree were collected, and FA indices were calculated. To examine differences in FA and herbivory within-individuals, we sampled pairs of mined and unmined leaves for asymmetry measurements. Differences in growth of leaf miners between leaf types were determined by tracing 50 mines of each species on symmetric leaves and asymmetric leaves. Asymmetric leaves contained significantly lower concentrations of tannins and higher concentrations of nitrogen than symmetric leaves for both plant species. Both frequency of asymmetric leaves on plants and levels of asymmetry positively influenced the abundance of Brachys, Stilbosis and other leaf miners, but no significant relationship between asymmetry and herbivory was observed for Acrocercops. Brachys and Stilbosis mines were smaller on asymmetric leaves, but differences in mine survivorship between symmetric and asymmetric leaves were observed only for Stilbosis mines. This study indicated that leaf miners might use leaf FA as a cue to plant quality, although differential survivorship among leaf types was not observed for all species studied. Reasons for the different results between guilds are discussed.     

Synchronous leaf production and herbivory in juveniles of Gustavia superba

Summary Synchronous leaf production has been proposed as a mechanism to reduce herbivore damage to young leaves by satiating herbivores. To test this hypothesis, I measured leaf production, leaf survivorship, and herbivore damage on juveniles of Gustavia superba (H.B.K.) Berg (Lecythidaceae), in two sites in Central Panama. Leaves were produced throughout the year, but there were peaks in leaf production at the beginning of the wet scason. Plants that produced leaves synchronously with conspecifics received significantly less damage than plants that produced leaves out of synchrony, and high levels of leaf damage were correlated with shorter leaf lifetimes. These data suggest that plant phenology can influence risks of herbivory.     

The apple sawfly, Hoplocampa testudinea: egg development and forecasting of egg hatch

The effect of temperature on egg development of the apple sawfly Hoplocampa testudinea Klug was studied under controlled conditions. Based on a linear model describing the relationship between temperature and developmental rates a thermal threshold of 6.9 °C and a thermal constant of 85 day-degrees were established. The experimental results were used to expand and parameterise an existing phenology model to include egg development and hatching of first instar larvae. The model thus became a valuable forecasting tool not only for adult emergence and the optimal monitoring period but also for egg hatch and the best timing of control measures against first instar larvae. The model was validated by comparing predictions with independent observations on egg-laying and subsequent hatching of first instar larvae under semi-field conditions.     

Effects of different leaf traits on growth rates of insect herbivores on willows

We examined relative effects of traits of leaf quality of ten willow species (Salix: Salicaceae) on growth rates of five species of insect herbivores found in interior Alaska (a willow sawfly, Nematus calais; the tiger swallowtail butterfly, Papilio canadensis; and three species of chrysomelid beetles, Gonioctena occidentalis, Calligrapha verrucosa, and Chrysomela falsa). Leaf traits examined were water content, toughness, total nitrogen contnet, pubescence, and presence or absence of phenolic glycosides. Of ten Salix species, four species contain phenolic glycosides in their leaves. We examined relative effects of water content, toughness, and nitrogen content of the Salix leaves on larval growth rates at three different levels, i.e., on a single host species, between different host species, and between herbivore species. The within-host analyses showed that effects of water content, toughness and/or nitrogen content on herbivore growth rates were generally significant in early-season herbivores but not in late-season herbivores. For each herbivore species, differences in growth rates between hosts were not explained by differences in water content, toughness, or nitrogen content. The between-herbivore analysis showed that the interspecific difference in larval growth rates were related to difference in water and nitrogen content of the hosts. Pubescence of Salix leaves had little effects on herbivore growth rates. Presence of phenolic glycosides had a positive effects on growth rates of a specialist, N. calais, but no effect on the other specialist, Ch. falsa. Presence of phenolic glycosides had, in general, negative effects on growth rates of nonspecialists, G. occidentalis, C. verrucosa, and P. canadensis.     

Population differences in host use by a seed-beetle: local adaptation, phenotypic plasticity and maternal effects

For insects that develop inside discrete hosts, both host size and host quality constrain offspring growth, influencing the evolution of body size and life history traits. Using a two-generation common garden experiment, we quantified the contribution of maternal and rearing hosts to differences in growth and life history traits between populations of the seed-feeding beetle Stator limbatus that use a large-seeded host, Acacia greggii, and a small-seeded host, Pseudosamanea guachapele. Populations differed genetically for all traits when beetles were raised in a common garden. Contrary to expectations from the local adaptation hypothesis, beetles from all populations were larger, developed faster and had higher survivorship when reared on seeds of A. greggii (the larger host), irrespective of their native host. We observed two host plant-mediated maternal effects: offspring matured sooner, regardless of their rearing host, when their mothers were reared on P. guachapele (this was not caused by an effect of rearing host on egg size), and females laid larger eggs on P. guachapele. This is the first study to document plasticity by S. limbatus in response to P. guachapele, suggesting that plasticity is an ancestral trait in S. limbatus that likely plays an important role in diet expansion. Although differences between populations in growth and life history traits are likely adaptations to their host plants, host-associated maternal effects, partly mediated by maternal egg size plasticity, influence growth and life history traits and likely play an important role in the evolution of the breadth of S. limbatus’ diet. More generally, phenotypic plasticity mediates the fitness consequences of using novel hosts, likely facilitating colonization of new hosts, but also buffering herbivores from selection post-colonization. Plasticity in response to novel versus normal hosts varied among our study populations such that disentangling the historical role of plasticity in mediating diet evolution requires the consideration of evolutionary history.     

Diversity of birch sawfly responses to seasonally atypical diets

Most insect herbivores are specialised on a particular plant taxon. To have a better understanding of host shift functions and consequences for insect herbivores, it is essential to gather more information on the effects of variation in host quality on specialists across species and environments. We examined the effects of seasonally atypical food on mortality, developmental time, and final body mass of six sawfly species (Hymenoptera: Symphyta) feeding on the foliage of mountain birch (Betula pubescens ssp. czerepanovii), whose pooled larval feeding periods form a gradient and cover the growing season. Insect phenology was manipulated so that the larvae of early-season species would feed on atypically mature leaves and mid- or late-season species would feed on atypically young leaves of their major host plant. Mortality increased dramatically for all species when the larval feeding schedule was advanced or delayed. This indicates a high degree of specialisation not only on a particular host but also to its phenological phases. The main cause of mortality on novel food was a rejection of the diet by the young larvae and their subsequent starvation. An interesting observation was that late-season species showed this response on nutritious young foliage. The effects of seasonally atypical diets on larval development and growth were species-specific and milder than the effects on mortality. Interestingly, for those individuals that accepted it, atypical food seemed to be most beneficial for species appearing at both ends of the seasonal gradient, which might be related to a wider exposure to variable food quality in natural conditions compared with other species. The diversity of responses to atypical food among closely related herbivore species with overlapping feeding periods on the same host plant is the most crucial finding of this study.     

Impact of epidermal leaf mining by the aspen leaf miner (Phyllocnistis populiella) on the growth, physiology, and leaf longevity of quaking aspen

The aspen leaf miner, Phyllocnistis populiella, feeds on the contents of epidermal cells on both top (adaxial) and bottom (abaxial) surfaces of quaking aspen leaves, leaving the photosynthetic tissue of the mesophyll intact. This type of feeding is taxonomically restricted to a small subset of leaf mining insects but can cause widespread plant damage during outbreaks. We studied the effect of epidermal mining on aspen growth and physiology during an outbreak of P. populiella in the boreal forest of interior Alaska. Experimental reduction of leaf miner density across two sites and 3 years significantly increased annual aspen growth rates relative to naturally mined controls. Leaf mining damage was negatively related to leaf longevity. Leaves with heavy mining damage abscised 4 weeks earlier, on average, than leaves with minimal mining damage. Mining damage to the top and bottom surfaces of leaves had different effects on physiology. Mining on the top surface of the leaf had no significant effect on photosynthesis or conductance and was unrelated to leaf stable C isotope ratio (delta(13)C). Mining damage to the bottom leaf surface, where stomata are located, had significant negative effects on net photosynthesis and water vapor conductance. Percent bottom mining was positively related to leaf delta(13)C. Taken together, the data suggest that the primary mechanism for the reduction of photosynthesis by epidermal leaf mining by P. populiella is the failure of stomata to open normally on bottom-mined leaves.     

Interspecific competition between the principal larval parasitoids of the pine sawfly,Neodiprion sertifer (Geoff.) (Hym.: Diprionidae)

Summary Interspecific competition between the 3 principal larval parasitoids of the pine sawfly, Neodiprion sertifer, is of common occurrence when total larval parasitism, and hence multiple attack rates, are high. At the intrinsic level, the ectoparasitoid Exenterus abruptorius is superior to the 2 endoparasitoids, Lophyroplectus luteator and Lamachus eques, respectively, whereas L. luteator is superior to L. eques only. During mass outbreaks of the host competition between Exenterus and Lophyroplectus is most intense, whilst Lamachus fails to build up in the presence of its 2 competitors (Fig. 1). However, due to its superior host finding ability, Lamachus dominates in areas of low host densities where the 2 other species are less efficient (Table 1). L. luteator frequently sustains heavy losses in direct competition with E. abruptorius, but these can be largely compensated in the next generation because its fertility is about 5-times higher than that of Exenterus.     

Early spring leaf out enhances growth and survival of saplings in a temperate deciduous forest

Saplings of many canopy tree species in winter deciduous forests receive the major portion of their light budget for their growing season prior to canopy closure in the spring. This period of high light may be critical for achieving a positive carbon (C) gain, thus contributing strongly to their growth and survival. This study of saplings of Aesculus glabra and Acer saccharum in Trelease Woods, Illinois, USA, tested this hypothesis experimentally by placing tents of shade cloth over saplings during their spring period of high light prior to canopy closure in three consecutive years. Leaf senescence began 16 days (year 0) and 60 days (year 1) earlier for shaded A. glabra saplings than control saplings. No change in senescence occurred for A. saccharum. The annual absolute growth in stem diameter of both species was negligible or negative for shaded saplings, but positive for control saplings. Only 7% of the shaded A. glabra saplings were alive after 2 years, while all control saplings survived for 3 years; only 20% of the shaded A. saccharum saplings survived for 3 years, while 73% of control saplings were alive after the same period. Early spring leaf out is a critical mechanism that allows the long-term persistence of saplings of these species in this winter deciduous forest. Studies and models of C gain, growth, and survival of saplings in deciduous forests may need to take into account their spring phenology because saplings of many species are actually “sun” individuals in the spring prior to their longer period in the summer shade.     

The apple sawfly, Hoplocampa testudinea: a temperature driven model for spring emergence of adults

The effect of temperature on post-diapause development of the apple sawfly Hoplocampa testudinea Klug was studied under controlled conditions. Survival was found to decrease at temperatures higher than 15°C, the lethal temperature being close to 25°C. The lower thermal threshold was 4.5°C; the values for males and females did not differ significantly. However, different values were established for unprotected individuals, in vermiculite and in potting compost respectively. Based on the overall mean and variance of the developmental time and the overall thermal threshold a simple soil temperature driven phenology model was constructed using a time- varying distributed delay. The validation with independent field and semi-field data revealed the model to be a reliable tool to predict apple sawfly phenology and hence to determine the optimal time for the installation of white sticky traps or for control measures such as insecticide treatments.     

Comparison of the physiology, morphology, and leaf demography of tropical saplings with different crown shapes

Branch architecture, leaf photosynthetic traits, and leaf demography were investigated in saplings of two woody species, Homolanthus caloneurus and Macaranga rostulata, co-occurring in the understory of a tropical mountain forest. M. rostulata saplings have cylindrical crowns, whereas H. caloneurus saplings have flat crowns. Saplings of the two species were found not to differ in area-based photosynthetic traits and in average light conditions in the understory of the studied site, but they do differ in internode length, leaf emergence rate, leaf lifespan, and total leaf area. Displayed leaf area of H. caloneurus saplings, which have the more rapid leaf emergence, was smaller than that of M. rostulata saplings, which have a longer leaf lifespan and larger total leaf area, although M. rostulata saplings showed a higher degree of leaf overlap. Short leaf lifespan and consequent small total leaf area would be linked to leaf overlap avoidance in the densely packed flat H. caloneurus crown. In contrast, M. rostulata saplings maintained a large total leaf area by producing leaves with a long leaf lifespan. In these understory saplings with a different crown architecture, we observed two contrasting adaptation strategies to shade which are achieved by adjusting a suite of morphological and leaf demographic characters. Each understory species has a suite of morphological traits and leaf demography specific to its architecture, thus attaining leaf overlap avoidance or large total leaf area.     

Enemy-free space? Host preference and larval performance of a willow leaf beetle

To examine whether enemy-free space is an important factor determining the host utilization pattern of a leaf beetle Plagiodera versicolora, we investigated the relationship between adult preference and offspring performance on three co-occurring willow species, Salix sachalinensis, S. miyabeana and S. integra. Salix sachalinensis was by far the most preferred host plant of feeding adults, while both S. miyabeana and S. integra were rarely fed upon. The fact that most oviposition was observed on S. sachalinensis also suggested that P. versicolora preferred S. sachalinensis to other willows for oviposition. This adult preference did not correspond well to patterns of larval performance on the three willow species in the absence of enemies. Higher survivorship, shorter developmental time and larger adult size were achieved on S. sachalinensis and S. miyabeana than on S. integra. Performance as indicated by female adult size and development time on S. miyabeana were higher than on S. sachalinensis. In the presence of enemies, however, the survivorship of first-instar larvae on S. miyabeana was much lower than on other willows. Adults of P. versicolora apparently avoided S. miyabeana as an oviposition and feeding host and preferred S. sachalinensis as an enemy-free space. This was not because larvae had poorer performance on S. miyabeana, but because predation pressure on eggs and early instar larvae was more severe on S. miyabeana.     

Effects of neem oil on feeding behaviour and development of the pear sawfly, Caliroa cerasi

Neem oil deterred feeding by pear sawfly, Caliroa cerasi L., larvae (Hymenoptera: Tenthredinidae), both in choice and in no-choice cherry leaf disk bioassays. Deterrence was greater in the choice tests, with 50% inhibition occurring at 0.49% aqueous neem oil compared with 1.11% in the no-choice tests. Antifeedant activity towards pear sawfly larvae is slightly less than has been observed for Lepidoptera, but is higher than deterrence to other insects such as aphids. Topical application of neem oil to sawfly larvae resulted in reduced feeding, increased mortality, and a trend towards slower development. Most larval mortality after neem treatment resulted from incomplete subsequent moults. The potential utilization of neem insecticides for control of pear sawfly in tree fruit pest management is discussed.     

Phenotypic selection on leaf water use efficiency and related ecophysiological traits for natural populations of desert sunflowers

Plant water-use efficiency (WUE) is expected to affect plant fitness and thus be under natural selection in arid habitats. Although many natural population studies have assessed plant WUE, only a few related WUE to fitness. The further determination of whether selection on WUE is direct or indirect through functionally related traits has yielded no consistent results. For natural populations of two desert annual sunflowers, Helianthus anomalus and H. deserticola, we used phenotypic selection analysis with vegetative biomass as the proxy for fitness to test (1) whether there was direct and indirect selection on WUE (carbon isotope ratio) and related traits (leaf N, area, succulence) and (2) whether direct selection was consistent with hypothesized drought/dehydration escape and avoidance strategies. There was direct selection for lower WUE in mesic and dry H. anomalus populations, consistent with dehydration escape, even though it is the longer lived of the two species. For mesic H. anomalus, direct selection favored lower WUE and higher N, suggesting that plants may be “wasting water” to increase N delivery via the transpiration stream. For the shorter lived H. deserticola in the direr habitat, there was indirect selection for lower WUE, inconsistent with drought escape. There was also direct selection for higher leaf N, succulence and leaf size. There was no direct selection for higher WUE consistent with dehydration avoidance in either species. Thus, in these natural populations of two desert dune species higher fitness was associated with some combination direct and indirect selection for lower WUE, higher leaf N and larger leaf size. Our understanding of the adaptive value of plant ecophysiological traits will benefit from further consideration of related traits such as leaf nitrogen and more tests in natural populations.     

Leaf-shape variation of Paederia foetida in Japan: reexamination of the small, narrow leaf form from Miyajima Island

Variations in Paederia foetida L. leaf shape were examined to evaluate the taxonomic validity of the small, narrow leaf form of P. foetida f. microphylla Honda from Miyajima Island, Honshu, Japan. There is considerable variation in P. foetida individuals in terms of leaf size and leaf index (leaf length:leaf width ratio). On Miyajima Island, some individuals have narrow leaves with a high leaf index value, a phenotype represented by the type specimen of P. foetida f. microphylla, and some do not. Given that the leaf size of individuals from Miyajima Island is smaller than that of individuals from other localities in Japan, and that the small leaf phenotype is stable even under cultivation, P. foetida f. microphylla is classified as the form having the smallest leaf size. Anatomical examination of leaf blades revealed that the large variation in leaf size was attributable to variation in the number of leaf cells but not to differences in cell size or cell shape. Based on these results, we discuss the endemism of P. foetida f. microphylla.     

Oviposition and larval performance of Rabdophaga terminalis on Salix spp. with special consideration to bud size of host plants

Females of the gall midge Rabdophaga terminalis (H. Loew) (Diptera: Cecidomyiidae) are very selective in their choice of host plants at oviposition. In preference trials, eggs were laid almost exclusively on S. alba and S. fragilis. One single oviposition occurred on S. viminalis but none on S. triandra, S. daphnoides or S. smithiana. Significantly more eggs were laid on S. alba than on S. fragilis. No differences were detected between clones of the same species regarding oviposition in laboratory trials. Within the species of S. alba, larger buds were preferred. However, this did not explain the preference of S. alba to S. fragilis. There was agreement between plant species selected by the females and survival of the larvae. No galls developed when eggs were transferred to species that were not oviposited on. There was no difference between S. alba and S. fragilis in larval growth, but after two weeks of development there was a significantly higher proportion of larvae in cocoons and pupae on S. alba. There was a negative linear relationship between weight of larvae and larval number per gall, indicating larval competition. The negative slope was steeper for S. alba than for S. fragilis when larvae were weighed after two weeks of development. There were some discrepancies between laboratory and field data on infestation of S. alba and S. fragilis. Higher infestation levels were related to denser stands in the field.

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Résumé Etude de la ponte et des performances larvaires de Rabdophaga terminalis (Diptera; Cecidomyiidae) sur plusieurs espèces de saules avec examen particulier de l'influence de la taille des bourgeons.La ponte et les performances larvaires de la cécidomyie galliforme, Rabdophaga terminalis (H. Loew) ont été étudiées sur plusieurs espèces et clones de saules cultivés en Suède. Les femelles sélectionnent avec soin les plantes hôtes lors de la ponte. Dans des essais préférentiels, les oeufs ont été pondus presque exclusivement sur S. alba et S. fragilis; une ponte a été observée sur S. viminalis et aucune sur S. triandra, S. daphnoïdes ou S. smithiana. Significativement plus d'oeufs ont été pondus sur S. alba que sur S. fragilis. Les essais au laboratoire n'ont pas permis de déceler des différences entre clones de la même espèce. Chez S. alba, les plus gros bourgeons étaient préférés, mais ceci ne permet pas d'expliquer la préférence pour S. alba par rapport à S. fragilis. Il y a coïncidence entre les plantes choisies par les femelles et al survie des larves; il n'y avait pas développement de galles quand des oeufs étaient transférés sur des espèces qui n'avaient pas reçu de pontes. La croissance larvaire sur S. alba et S. fragilis ne présentait pas de différences, mais après 2 semaines de développement il y avait significativement plus de larves en cocons et pupes sur S. alba. La corrélation entre le poids des larves et le nombre de larves par cellule était linéaire et négative, ce qui indique une compétition larvaire; la pente était plus raide pour S. alba que pour S. fragilis quand les larves étaient pesées au bout de 2 semaines de développement. Il y a eu quelques discordances entre les résultats au laboratoire et les données de terrain sur la contamination de S. alba et S. fragilis. Les plus fortes contaminations étaient observées à l'extérieur sur pépinières denses.