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

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

Feeding of pea leafminer larvae simultaneously activates jasmonic and salicylic acid pathways in plants to release a terpenoid for indirect defense

The pea leafminer, Liriomyza huidobrensis, is an important pest species affecting ornamental crops worldwide. Plant damage consists of oviposition and feeding punctures created by female adult flies as well as larva-bored mines in leaf mesophyll tissues. How plants indirectly defend themselves from these two types of leafminer damage has not been sufficiently investigated. In this study, we compared the indirect defense responses of bean plants infested by either female adults or larvae. Puncturing of leaves by adults released green leaf volatiles and terpenoids, while larval feeding caused plants to additionally emit methyl salicylate and (E,E)-4,8,12-trimethyl-1,3,7,11-tridecatetraene (TMTT). Puncturing of plants by female adults induced increases in jasmonic acid (JA) and JA-related gene expressions but reduced the expressions of salicylic acid (SA)-related genes. In contrast, JA and SA and their-related gene expression levels were increased significantly by larval feeding. The exogenous application of JA+SA significantly triggered TMTT emission, thereby significantly inducing the orientation behavior of parasitoids. Our study has confirmed that larval feeding can trigger TMTT emission through the activation of both JA and SA pathways to attract parasitoids; however, TMTT alone is less attractive than the complete blend of volatiles released by infested plants.     

Nutritional quality of specific leaf tissues and selective feeding by a specialist leafminer

Summary Many folivorous insects are selective feeders which consume specific leaf tissues. For specialist herbivores feeding on plants of overall low nutritional quality, selective feeding may allow consumption of a high quality resource. Selective feeding may also allow insects to avoid structural or allelochemical defenses. We examined the structure and chemistry of leaves of American holly, Ilexopaca Aiton, and the feeding site of its principal insect herbivore, the native holly leafminer, Phytomyza ilicicola Loew (Diptera: Agromyzidae), to test the hypothesis that the leafminer consumes tissues which are of greater nutritional quality than the leaf as a whole. Holly leaves have a continuous layer of palisade mesophyll, uninterrupted by fibers or vascular bundles. The leafminer feeds entirely within this layer. The palisade mesophyll contained 196 mg/g dry wt extractable protein, more than twice as much as the leaf as a whole, and 375 mg/g dry wt saponins, more than 9 times that of the leaf as a whole. The water content of the palisade mesophyll was 66% higher than that of the leaf as a whole. The palisade mesophyll is 3–4 cell layers thick in leaves grown in full sun, but only 2 layers thick in shaded leaves. Crystals, probably of calcium oxalate, are abundant in the abaxial cell layer. These may impose mechanical constraints on larval feeding in shade leaves, which are thinner than sun leaves. Selective feeding on the middle palisade mesophyll of sun leaves allows the leafminer to consume a resource which is lacking in mechanical barriers and is rich in protein and water, but which contains large amounts of saponins.The investigation reported in this paper (No. 86-8-7-117) is in connection with a project of the Kentucky Agricultural Experiment Station and is published with the approval of the Director     

Oviposition site selection by a lepidopteran leafminer in response to heterogeneity of leaf surface conditions: structural traits and microclimates

1. Leafminer larvae are sedentary and make feeding tracks called mines. Their spatial distribution in trees affects their growth and survival through interaction with the heterogeneity of environments, such as leaf traits and microclimate. Lepidopteran leafminers that mine lower leaf surfaces have shown evolutionary radiation, suggesting that lower surfaces improve leafminer performance. 2. The lepidopteran multivoltine leafminer Phyllocnistis sp. Zeller (Gracirallidae: Phyllocnistinae) uses the Japanese privet Ligustrum japonicum Thunb. (Oleaceae). It mines only the lower‐surface epidermal layer of primary shoot leaves early in the occurrence season, but once lammas shoots appear, which happens in seasons other than spring, it preferentially uses the lower surface, but also uses the upper surface of the leaves. This study examined whether selection of oviposition sites was associated with the structural traits and microclimate of the leaf surface. 3. The shift of oviposition site from primary to lammas shoot leaves followed increasing hardness and epidermal cell wall thickness of primary shoot leaves during leaf development, and mine initiation rates decreased below 20% after oviposition on mature primary shoot leaves. Preference for the lower surface was related to the thinner cuticle. However, the thinner cuticle of the upper surface on lammas shoot leaves allowed Phyllocnistis sp. to expand its mining sites to both surfaces with no decrease in mine initiation and emergence rates. 4. Microclimates (leaf surface and mine temperatures) did not differ between upper and lower surfaces, suggesting that microclimate did not affect oviposition site selection by Phyllocnistis sp. These results suggest that the adaptive radiation of lower‐surface mining may have been influenced by the leaf surface characteristics.     

Trichome-borne and artificially applied acylsugars of wild tomato deter feeding and oviposition of the leafminer Liriomyza trifolii

Oviposition and adult feeding of the leafminer Liriomyza trifollii (Burgess) (Diptera, Agromyzidae) on Lycopersicon pennellii (Corr.) D'Arcy and its F₁ hybrid with Lycopersicon esculentum (Mill.) was significantly less than that on the cultivated tomato, L. esculentum. The resistance of L. pennellii and the F₁ was reduced following rinsing of foliage with ethanol. Resistant attributes of L. pennellii were transferred to L. esculentum through appression of L. pennellii foliage to L. esculentum leaflets. Application of purified 2,3,4-tri-O-acylglucoses (the principal component of type IV glandular trichome exudate of L. pennellii) to L. esculentum significantly decreased feeding and oviposition on L. esculentum leaflets by 61–99%. Therefore the principal mechanism of resistance to this leafminer by L. pennellii is the secretion of these acylglucoses. Dose response analysis of acylglucoses applied to L. esculentum shows that dosages as low as 10% those found on L. pennellii provide large reductions (91%) in leaf punctures and mines.     

Adult movement of the native holly leafminer,Phytomyza ilicicola Loew (Diptera: Agromyzidae): consequences for host choice within and between habitats

Summary Phytomyza ilicicola is a specialist herbivore, the larvae of which mine, and the adults of which feed and oviposit on the developing leaves of American Holly, Ilex opaca. Adult Phytomyza have been shown to discriminate among host plants on the basis of several host and habitat factors. Such discrimination may explain, in part, the observed density variation of larval Phytomyza among individual trees and between forest and suburban habitats. In order for discrimination to influence density, adult Phytomyza must be sufficiently vagile to move among habitats and to encounter many hosts. We experimentally monitored the movement of adults to ask if this were so. In addition we asked whether inter-host movements were altered by resource concentration, and whether vagility offered an escape from parasitism. To answer these questions, three isolated holly trees to serve as colonization sources were selected in early spring, prior to adult emergence. Potted target trees were then placed singly and in groups of four at 3 compass directions around and at 3 different distances (10, 25, and 50 m) from each of the source trees. Six weeks after exposure to adult Phytomyza, the number of feeding punctures and first instar larvae were counted. Clones were then overwintered in cold frames and in early spring, just prior to adult emergence, all pupae were examined for parasitism by the dominant parasitoid Opius striatriventris. Feeding and parasitism were similar at all 3 distances from the source trees, indicating that Opius was at least as vagile as Phytomyza. However, first instar larvae decreased with distance. There was no evidence that resource concentration affected adult movement. Our results suggest that Phytomyza are sufficiently vagile to choose among hosts within and between habitats. However, movement does not translate into equally high oviposition at all distances from a source, nor does it provide an escape from parasitism by Opius.     

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.     

Growth responses of an African savanna tree, <Emphasis Type="Italic">Bauhinia thonningii</Emphasis> Schumacher,to defoliation,fire and climate

This study investigated the growth responses to defoliation, fire-exposure and climate factors of a widespread Africana savanna tree, Bauhinia thonningii Schumacher, at a site in central Zambia. Experimental trees (n = 47) were either exposed to fire (n = 12) in the first half of the dry season or protected from fire (n = 35). Some of the fire-protected trees (n = 12) were subjected to artificial defoliation in two consecutive years. Phenological responses (bud break, leaf flush and leaf production) to fire-exposure and defoliation were monitored on permanently marked sample shoots over a 2-year period. Radial tree growth (diameter at 1.3 m above ground) was measured annually over a 7-year period from 1998 to 2005. During the first two years, defoliation and fire-exposure advanced the onset of bud break and leaf flush but fire-exposed trees produced significantly less leaves than did trees protected from fire, probably because scorching caused more severe shoot die-back than is normal. Leaf production was also significantly affected by experimental treatments and their interaction with year. Although treatments had significant short-term effects on radial growth, previous-year growth significantly influenced current-year growth, thereby confirming the existence of autocorrelation in the time-series growth data of B. thonningii. The interaction between previous-year growth and climate factors explained a significant proportion (25–40%, P < 0.001) of the variance in annual tree growth. Variogram models predicted that a 2-year manual defoliation treatment would shorten the longer-term growth cycle while continuous fire-exposure extended the cycle by one year. The results are useful for the management of savanna trees.     

Variability of Lycopersicon hirsutum f. typicum and possible compounds involved in its resistance to Tuta absoluta

1 The resistance of Lycopersicon hirsutum f. typicum and L. esculentum to the leafminer Tuta absoluta (Meyrick) (Lepidoptera: Gelechiidae) was investigated in two experiments. The first experiment assessed the resistance of L. hirsutum f. typicum (accession LA 1777) to T. absoluta, the second assessed the variability of the resistance among plants of this accession. 2 It was found that L. hirsutum f. typicum was resistant to the leafminer, which showed longer larval phase, higher larval mortality, smaller numbers of large mines and greater numbers of small mines/leaf than on L. esculentum. Seven plants of the accession LA 1777 showed lower number of small mines/leaf, but their results of larval mortality and length of larval and pupal phase were similar. Twenty-two peaks were observed in total ion chromatograms of L. hirsutum f. typicum hexane extracts of leaves subjected to a gas chromatographic-mass spectrometric analysis (CG-MS). However, the matches obtained between the peak spectra and the MS database were too low to justify even preliminary identification of the compounds.     

Nonglandular leaf trichomes as short-term inducible defense of the grey alder,Alnus incana (L.), against the chrysomelid beetle,Agelastica alni L.

Summary The grey alder compensates leaf area losses due to insect grazing by continuously producing new leaves throughout the vegetative period. Different degrees of defoliation were attained experimentally by a controlled release of the oligophagous beetle Agelastica alni on arbitrarily selected trees from a homogenous population of young alders. The reduction in leaf area per tree significantly influenced the density of leaf trichomes, assessed 10–30 days later, on newly sprouting leaves only. Cross-correlations between leaf area reduction and trichome density were strongest for leaves which completed unfolding 14–21 days after damage. Dualchoice assays suggested a negative influence of trichomes on oviposition rate of A. alni. Removal of trichomes by shaving demonstrated the highly significant effect of trichomes on feeding behavior of adults and larvae in dual-choice assays. The role of the induced increase in trichome density as a possible short-term defense reaction against herbivorous insects is discussed.     

Plant phenology-mediated indirect effects: The gall midge opens the phenological window wider for a leaf beetle

We examined whether larvae of the gall midge Rabdophaga rigidae (Diptera: Cecidomyiidae) can modify the seasonal dynamics of the density of a leaf beetle, Plagiodera versicolora (Coleoptera: Chrysomelidae), by modifying the leaf flushing phenology of its host willow species, Salix serissaefolia and Salix eriocarpa (Salicaceae). To test this, we conducted field observations and a laboratory experiment. The field observations demonstrated that the leaf flushing phenology of the willows and the seasonal dynamics of the beetle density differed between shoots with stem galls and shoots without them. On galled shoots of both willow species, secondary shoot growth and secondary leaf production were promoted; consequently, leaf production showed a bimodal pattern and leaf production periods were 1 to 2 months longer than on non‐galled shoots. The adult beetle density on galled shoots was thus enhanced late in the season, and was found to change seasonally, synchronizing with the production of new leaves on the host willow species. From the results of our laboratory experiment, we attributed this synchrony between adult beetle density and willow leaf flush to beetles’ preference to eat new leaves rather than old. Indeed, beetles consumed five times more of the young leaves when they were fed both young and old leaves. These results indicate that stem galls indirectly enhance the adult beetle density by enhancing food quality and quantity late in the beetle‐feeding season. We therefore conclude that midge galls widen the phenological window for leaf beetles by extending the willows’ leaf flush periods.     

Interactive effects of leaf maturation and phenolics on consumption and growth of a geometrid moth

Phenolic compounds are commonly regarded as the main chemical defenses of deciduous woody plants against insects. To examine how indices of leaf maturation (water content, toughness, and sugar/protein ratio) modified larval consumption and growth relative to phenolics and phenolic-related leaf traits, we measured consumption and growth of fourth-instar Epirrita autumnata (Bkh.) (Lepidoptera: Geometridae) larvae on three different days on young, normal, and mature leaves, respectively, from the same mountain birch (Betula pubescens ssp. czerepanovii (Orlova) Hämet-Ahti) trees. The larvae achieved the same growth rates on young and normal leaves, but had to consume 40% more on the latter. On more mature leaves, larval growth was poorer and was positively correlated with sugar/protein ratios (although the ratio peaked at that time). Indices of leaf maturation correlated with several phenolics in data pooled over the three study days, but poorly in any individual day. Similarly, in the pooled data, larval consumption and growth correlated with several leaf traits, but correlations between leaf and insect traits were few on any of the three days, and no trait was significant on each of the three days.We next examined whether variation in the maturation indices modified the associations of phenolics with insect consumption and growth. When interactions between phenolics and leaf maturation indices were taken into account, the number of phenolic compounds displaying significant associations with insect traits more than doubled. The relative importance of interactive versus direct associations increased with leaf maturation: on young leaves five phenolics showed direct and eleven interactive associations with insect traits, while in mature leaves we found two phenolics to display direct and thirteen phenolics interactive associations. Leaf water content, either alone or together with toughness and sugar/protein ratio, generally explained more of the variance in Epirrita growth (up to 59%) than any phenolic or phenolic-related trait alone (highest value 20%). Including interactive effects between phenolics and indices of leaf maturation in the model increased the proportion explained of variance in larval growth between 49 and 73%. Maturation indices explained 0 to 23% of variance in consumption, and the phenolic compound with the highest (positive!) correlation alone up to 28%, but taking into account interactions between phenolics and maturation indices raised the degree of explanation much (namely, 32 to 53%) over that explained by indices of leaf maturation alone. This indicates strong interactive effects on consumption between phenolics and indices of leaf maturation.     

Anatomical Injury Induced by the Eriophyid Mite Aceria anthocoptes on the Leaves of Cirsium arvense

Anatomical injury of the leaves of the invasive species, Cirsium arvense (L.) Scop., caused by the eriophyid mite Aceria anthocoptes (Nal.), which is the only eriophyid mite that has been recorded on C. arvense worldwide, is described. The injury induced by the mite feeding on the leaves of C. arvense results in visible russeting and bronzing of the leaves. Other conspicuous deformations are folding and distortion of the leaf blade and curling of leaf edge, as well as gradual drying of leaves. The anatomical injury of the mature leaves of field-collected plants was limited to the epidermis of the lower leaf surface. However, on young leaves of experimentally infested plants, rust mite injuries extend to epidermal cells on both leaf surfaces and to those of deeper mesophyll layers. On these leaves, lesions on the lower leaf surface even affected the phloem of the vascular bundles. Leaf damage induced by A. anthocoptes is discussed with regard to the mite’s potential as a biological control agent of C. arvense.     

Leaf mines as visual defensive signals to herbivores

Leaf‐mining insects produce conspicuous and distinct leaf mines on various types of plant leaves. The diversity of leaf‐mine morphology has typically been explained by several factors, such as selective feeding on plant tissues, improvement of microclimate, faecal disposal, reduction in the efficiency of parasitoid search behaviour and leafminer phylogeny. Although these factors are certainly associated with mining patterns, masking the mines, rather than making them conspicuous, appears to be more advantageous for deterring parasitoids and predators of leafminers. However, here, I propose that prominent leaf mines may serve to signal or cue herbivores to avoid feeding on the mined leaves. Because most leafminers are sessile and complete their development within a single leaf, herbivory of mined leaves is detrimental to leafminer survival. Other herbivores appear to avoid consuming mined leaves for a variety of reasons: leaf mines mimic leaf variegation or mottling; mined leaves induce chemical and physical defences against herbivores; and leaf mines mimic fungal infection, animal excrement, and necrosed plant tissues. Hence, natural selection may have favoured leafminers that produce conspicuous mines because of the increased survival and fecundity of thereby reducing herbivory on mined leaves.     

Biosynthesis and metabolism of purine alkaloids in leaves of cocoa tea (Camellia ptilophylla)

The biosynthesis and metabolism of purine alkaloids in leaves ofCamellia ptilophylla (cocoa tea), a new tea resource in China, have been investigated. The major purine alkaloid was theobromine, with theophylline also being present as a minor component. Caffeine was not accumulated in detectable quantities. Theobromine was synthesized from [8-¹⁴C] adenine and the rate of its biosynthesis in the segments from young and mature leaves from flush shoots was approximately 10 times higher than that from aged leaves from 1-year old shoots. Neither cellfree extracts nor segments fromC. ptilophylla leaves could convert theobromine to caffeine. A large quantity of [2-¹⁴C] xanthine taken up by the leaf segments was degraded to¹⁴CO₂ via the conventional purine catabolic pathway that includes allantoin as an intermediate. However, small amounts of [2-¹⁴C] xanthine were also converted to theobromine. Considerable amounts of [8-¹⁴C] caffeine exogenously supplied to the leaf segments ofC. ptilophylla was changed to theobromine. These results indicate that leaves ofC. ptilophylla exhibit unusual purine alkaloid metabolism as i) they have the capacity to synthesize theobromine from adenine nucleotides, but they lack adequate methyltransferase activity to convert of theobromine to caffeine in detectable quantities, ii) the leaves have a capacity to convert xanthine to theobromine, probably via 3-methylxanthine.     

Sources of variation in rapidly inducible responses to leaf damage in the mountain birch-insect herbivore system

Summary Studies on rapidly inducible resistance in trees against insect herbivores show substantial variation in the strength of responses. Here we report the results of a study which examined causes of this variation. We bioassayed the quality of leaves of two developmental phases (young vs. mature) of the mountain birch Betula pubescens ssp. tortuosa by measuring the growth of two instars of Epirrita autumnata larvae. We used only short shoot leaves from trees of a natural stand, uniform in size and age. Damage was caused by larvae and artificial tearing of leaf lamina, varying the scale and time. We separated seasonal changes in plants from instar-dependent effects of the animals by testing experimental larvae in two subsequent growth trials. We found that only larval-made damage induced responses in leaves that made the leaves significantly poorer quality for the test larvae. Artificial damage induced only weak responses, and artificial canopy-wide damage even caused slight improvement of leaf quality. Cumulative leaf damage did not strengthen birch responses. Leaves that were in the expansion phase responded to damage while fully-expanded, mature leaves showed no response. The pattern of responses indicated that there might be physiological constraints: small-scale damage induced resistance against the larvae but largescale damage did not. Prevalent weather conditions might have modified these responses. Larvae of two instars and sexes, of low- and high-density populations responded to leaf damage similarly. However, prior experience of larvae with the host plant may have affected subsequent larval performance. Variation in rapidly inducible responses in birches was caused by plant characters rather than by test animals.     

Effects of plant chemical extracts and physical characteristics of Apium graveolens and Chenopodium murale on host choice by Spodoptera exigua larvae

Choice tests with whole plants and leaf discs indicated that fourth instar Spodoptera exigua (Hübner) (Noctuidae: Amphypyrini) were found more frequently and ate significantly more of the weed Chenopodium murale than the associated crop plant Apium graveolens. In order to explain the preference, plant extracts, plant volatiles, soluble protein concentrations, water contents, and leaf toughness of the two plants were investigated. Bioassays of aqueous methanol (90%) and hexane extracts of leaves on cellulose discs indicated that neither attractants in C. murale nor repellents in A. graveolens could account for the observed preference. No significant difference could be found between the effects of plant volatiles from C. murale, A. graveolens and a control on larval dispersal by S. exigua. Selective feeding for higher levels of proteins also was not a factor, because A. graveolens had nearly twice the soluble protein of C. murale. Water content was approximately 6% higher (by weight) in C. murale than A. graveolens but most polyphagous larvae do not typically show compensatory feeding for water alone. However, the potentially related characteristic of leaf toughness was significantly different, with A. graveolens exhibiting 1.53 times the toughness of C. murale. Studies comparing five types of larval behavior on both plant species showed that the time spent in swallowing behavior was significantly greater on the tougher A. graveolens leaves relative to C. murale. To test the hypothesis that leaf toughness was affecting larval host choice, both plants were finely ground and incorporated into agar blocks. No differences in feeding behavior were detected. The implications of leaf toughness for larval diet and host choice are discussed.     

Variation in shoot mortality within crowns of severely defoliated <Emphasis Type="Italic">Betula maximowicziana</Emphasis> trees in Hokkaido,northern Japan

To clarify mortality patterns of current-year shoots within the crown of Betula maximowicziana Regel after severe insect herbivory in central Hokkaido, northern Japan, we investigated the degree of defoliation, pattern of shoot development, shoot mortality, and leaf tissue-water relations. One hundred current-year long shoots growing in a B. maximowicziana plantation were observed for defoliation and mortality in June 2002. An outbreak of herbivorous insects (Caligula japonica and Lymantria dispar praeterea) occurred in the stand in mid-to-late June, and the monitored shoots were defoliated to various degrees. Within 1 month of defoliation, some of the severely defoliated shoots had produced new leaves on short shoots that had emerged from axillary buds. Stepwise logistic regression revealed that the probability that current-year long shoots would put out axillary short shoots with leaves is closely related to the degree of defoliation. To evaluate the water relations of the leaves, we determined pressure–volume curves for the leaves that survived the herbivorous insect outbreak and the new leaves that emerged after defoliation. The water potential at turgor loss (Ψ_l,tlp) and the osmotic potential at full turgidity (Ψ_π,sat) were higher for the new leaves than for the surviving leaves, indicating a lower ability to maintain leaf cell turgor against leaf dehydration in the new leaves. Of the 100 shoots, 13 died after the emergence of new leaves. Stepwise logistic regression revealed that the probability that the long shoots would die generally increased with the emergence of new leaves, with increasing shoot height. This result suggests that the combined effect of the vulnerability of newly emerged leaves and low water availability, associated with higher shoot positions within the crown, caused shoot mortality. Based on our results, some possible mechanisms for mortality in severely defoliated B. maximowicziana are discussed.     

Growth of coyote willow and the attack and survival of a mid-rib galling sawfly,Euura sp.

We studied the relationship between variation in age and shoot characteristics of the host plant Salix exigua Nuttall (coyote or sandbar willow) and the attack and survival of Euura sp. (an unnamed leaf-midrib galling sawfly). Variation in shoot characteristics resulted from reduced growth as willow ramets aged. Mean shoot length per ramet and mean longest leaf length per shoot decreased by 95% and 50% respectively between 1- and 9-year-old willow ramets. All measured shoot characteristics-shoot length, longest leaf length, number of leaves per shoot, and mean internode length-were significantly negatively correlated with ramet age (r ² ranged from –0.23 to –0.41). Correlations between shoot characteristics were highly positive, indicating that plants also grew in a strongly integrated fashion (r ² ranged from 0.54 to 0.85). Four hypotheses were examined to explain sawfly attack patterns. The host-plant hypothesis was supported in explaining enhanced larval sawfly survival through reduced plant resistance. As willow ramets aged, the probability of Euura sp. attack decreased over 10-fold, from 0.315 on 1-year-old ramets to 0.024 on 2- to 9-year-old ramets. As shoot length increased, the probability of sawfly attack increased over 100-fold, from 0.007 on shoots <100 mm, to 0.800 on shoots in the 1001–1100 mm shoot length class. These attack patterns occurred even though 1-year-old ramets and shoots >500 mm each represented less than 2% of the total shoots available for oviposition. Host plant induced mortality of the egg/early instar stage decreased by 50% on longer leaves and was the most important factor determining survival differences between vigorous and non-vigorous hosts. Sawfly attack was not determined by the resource distribution hypothesis. Although shoots <200 mm contained 82% of the total leaves available, they contained only 43% of the galls initiated. The attack pattern also was not explained by the gall volume hypothesis. Although gall volume increased on longer shoots, there was no significant variation in mid or late instar mortality over shoot length, as would be expected if food resources within smaller galls were limited. The natural enemy attack hypothesis could not explain the pattern of oviposition since predation was greater on longer shoots and leaves. In addition, larval survival was related to oviposition behavior. Due to a 69% reduction in late instar death and an 83% reduction in parasitism, survival of progeny in galls initiated close to the petiole base was 2.8 times greater than in galls initiated near the leaf tip. A 75% reduction in gall volume over this range of gall positions may account for the observed increases in late instar mortality and parasitism.     

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.