Hormaphis hamamelidis and gall size: a test of the plant vigor hypothesis

We tested the Plant Vigor Hypothesis by determining the distribution of galls formed on leaves of witch hazel, Hamamelis virginiana , by the aphid Hormaphis hamamelidis , and by determining various factors that affect the fecundity of the gall-forming fundatrices. We also studied the role of the fundatrix in host plant manipulation. While the mean number of galls per leaf was low, galls had an aggregated distribution among leaves. Among trees, the average number of galls per leaf was not related to the mean leaf size, contrary to the preference prediction of the Plant Vigor Hypothesis. While fundatrices preferred the distal leaves of buds, which grew more than the proximal leaves, being on distal leaves conferred no increase in fecundity for fundatrices, contrary to predictions of the Plant Vigor Hypothesis. Gall size was the factor that explained the largest proportion of variation in fundatrix fecundity; fundatrix size explained somewhat less of the variation. Also, gall position on the leaf, number of aphid galls on the leaf, and on which leaf of the bud the gall was located all played small, statistically significant roles in explaining fundatrix fecundity, but their effects were variable between experiments. Removal of fundatrices shortly after galls had enclosed them limited the growth of galls, indicating the role of the fundatrices in gall growth. We compare and contrast this system versus other gall-forming insects, as well as discuss the adaptive significance of the aphid manipulation of the host plant. Much of the data contradict predictions of the Plant Vigor Hypothesis, and we discuss how gall size, as a measure of plant growth caused by insect manipulation, explains the observed patterns.     

Mechanisms of hydraulic conductivity in the leaf galls of Meunieriella sp. (Cecidomyiidae) in Avicennia schaueriana (Acanthaceae): does vascularization explain the preferred sites of induction?

• Intralaminar galls of Meunieriella result from ground tissue proliferation in leaves of Avicennia schaueriana, a typical halophytic mangrove. We tested if the preferred sites of gall induction were the midribs and secondary veins (SV) at the basal leaf portion, where the galls were expected to be largest; and if the vascular system in galls and adjacent regions was altered to favour water supply in galls, thus increasing their growth.
• Gall induction sites and gall sizes were quantified according to leaf portions and regions. Anatomical and histometric analyses in vascular and ground tissues of galls and adjacent regions were compared to equivalent regions of non-galled leaves.
• The galls were largest at basal sites on leaves, the midrib and SV. More galls occurred on the apical portion of the leaf, and on the leaf blade and secondary vein regions. Changes in shape and vascular system area, number and diameter of vessel elements were detected in both galls and adjacent regions. Fewer and smaller-sized vessel elements were observed in regions proximal to the galls and inside them.
• Gall size is not related with preferred induction sites, which could be explained by factors such as thermal balance. Alterations in the vascular system indicate reduced hydraulic conductivity in the xylem in the proximal region and inside galls, leading to water leakage to gall parenchyma cells. This compensatory mechanism explains the expansion and proliferation of water storage and spongy parenchyma cells in the galls, explaining the higher growth in more vascularized regions.

     

The effect of hosptlant and parasitoids on the reproductive success of the parthenogenetic gall wasp Diplolepis rosae (Hymenoptera,Cynipidae)

Summary The univoltine cynipid gall wasp Diplolepis rosae reproduces by an obligate homozygosity promoting system known as gamete duplication. The wasp is confined to roses (Rosa spp) on which it induces large, complex and multichambered galls. In southern Sweden, D. rosae was found to parasitize Rosa canina, R. dumalis, R. rubiginosa, R. villosa, R. sherardi and R. rubrifolia, but not R. majalis and R. rugosa. The distribution of galls shows that there are differences in the relation between wasp and hosplant with respect both to species and individual plants. There is a positive correlation between wasp size and gall (clutch) size. Parasitoid pressure was found to be high, causing D. rosae an estimated average larval loss of approximately 75%, mainly due to the attack of the ichneumonid wasp Orthopelma mediator. The very common cynipid inquiline Periclistus brandtii does not seem to have any negative effects. Overall parasitism and probability of no hatched offspring per gall decrease with increasing gall (clutch) size. The probability of loosing all of a given number of offspring decreases with the number of galls produced. It is suggested that D. rosae, in order to escape parasitoids, needs high ability to establish new colonies. Hence the production of many comparatively small galls, which increases the chance of leaving any offspring, rather than the production of few large galls, maximizing the number of offspring, should be favoured by selection.     

Intra‐plant heterogeneity influences the preference and performance of juveniles and adults of a gall midge

Abstract 1. Field studies were conducted to evaluate the preference and performance of a gall‐inducing midge (Harmandia tremulae) within the crown of trembling aspen (Populus tremuloides). Females did not select oviposition sites preferentially within leaves, but did lay preferentially on young leaves. 2. Larvae were the only life stage involved in gall site selection within leaves and in gall initiation and development. Gall size, which was positively related to survival, was highest for galls on mid veins that were located close to the petiole. However, one‐third of galls were located on lateral veins and most galls were not adjacent to the petiole, indicating that many larvae choose sub‐optimal gall initiation sites. 3. Gall density was positively associated with leaf length, and leaf length, was positively associated with gall size. However, gall density per leaf was not related to larval survival in galls. This latter result may be a result of an observed inverse relationship between gall size and gall density for similar‐sized leaves. 4. The results partially support the plant vigour and optimal plant module size hypotheses, which predict that galler fitness in successfully induced galls should be highest on large, fast‐growing plant modules. The lack of a strong preference‐performance link supports the confusion hypothesis, which predicts that oviposition and gall site selection may often be suboptimal in systems where galler lifespan is short. This study suggests that small‐scale variations in plant quality within leaves, can render gall site selection by juveniles as important as that previously reported for adult females.     

Trade‐off between fecundity and survival generates stabilizing selection on gall size

Complex interactions within multitrophic communities are fundamental to the evolution of individual species that reside within them. One common outcome of species interactions are fitness trade‐offs, where traits adaptive in some circumstances are maladaptive in others. Here, we identify a fitness trade‐off between fecundity and survival in the cynipid wasp Callirhytis quercusbatatoides that induces multichambered galls on the stem of its host plant Quercus virginiana. We first quantified this trade‐off in natural populations by documenting two relationships: a positive association between the trait gall size and fecundity, as larger galls contain more offspring, and a negative association between gall size and survival, as larger galls are attacked by birds at a higher rate. Next, we performed a field‐based experimental evolution study where birds were excluded from the entire canopy of 11 large host trees for five years. As a result of the five‐year release from avian predators, we observed a significant shift to larger galls per tree. Overall, our study demonstrates how two opposing forces of selection can generate stabilizing selection on a critical phenotypic trait in wild populations, and how traits can evolve rapidly in the predicted direction when conditions change.     

A nectar-secreting gall wasp and ant mutualism: selection and counter-selection shaping gall wasp phenology, fecundity and persistence

Abstract.

• 1 The natural history of a gall wasp including interactions with inquilines, parasites, and a mutualistic ant are examined. The stability of the system is described from the perspective of influences on gall wasp life history characteristics.
• 2 An exclusion experiment demonstrated that the nectar-secreting gall of Disholcaspis perniciosa mediates a mutualism with the tending ant, Formica obscuripes. Survivorship increased from 0% in the absence of ants to 25.3% in their presence, largely due to the exclusion of inquilines.
• 3 Specialized parasites, Eudecatoma spp., attacked before the ant-gall interaction began, when the developing gall was still beneath the host plant (Quercus gambellii) epidermis and ants were not in attendance. They may select for later developing gall wasps, which benefit by having fewer individuals parasitized. However, counter-selection for earlier development may result from decreased gall wasp size, decreased fecundity, and an increase in gall failures resulting from late development.
• 4 Local persistence of the gall wasp population despite increased pressure from inquilines and parasites was attributed to gall wasp escape in time due to polymorphic emergence resulting from diapause. Most individuals emerge at the end of the summer, but approximately 15% remain in the galls as prepupae for 1–5 years.

     

Influence of galls of Phanacis taraxaci on carbon partitioning within common dandelion, Taraxacum officinale

We examined how leaf galls, induced by the cynipid wasp Phanacis taraxaci, influence the partitioning of photoassimilates within the host, the common dandelion, Taraxacum officinale. Galled and ungalled plants were exposed to ¹⁴CO₂ and the labelled photoassimilates accumulating within galls and other parts of the host were measured. During the growth phase of the gall they were physiological sinks for photoassimilates, accumulating 9% to 70% of total carbon produced by the host, depending upon the number of galls per plant. High levels of ¹⁴C assimilation in the leaves of galled plants compared to controls, suggest that galls actively redirect carbon resources from unattacked leaves of their host plant. This represents a significant drain on the carbon resources of the host, which increases with the number and size of galls per plant. Active assimilation of ¹⁴C by the gall is greatest in the growth phase and is several orders of magnitude lower in the maturation phase. This finding is consistent with physiological and anatomical changes that occur during the two phases of gall development and represents a key developmental strategy by cynipids to ensure adequate food resources before larval growth begins.     

The influence of pruning on wasp inhabitants of galls induced by Hemadas nubilipennis Ashmead (Hymenoptera: Pteromalidae) on lowbush blueberry

The efficacy of pruning methods for managing blueberry stem galls caused by the chalcid wasp, Hemadas nubilipennis (Ashmead), was studied in five commercial lowbush blueberry (Vaccinium angustifolium Aiton) fields in Nova Scotia, Canada, between October 1999 and May 2000. Blueberry fields were mowed in the fall, and burning treatments were subsequently applied either in the fall or the spring. Three treatments were compared: mowing only, mowing plus fall burning, and mowing plus spring burning. Galls collected from the mow plus spring-burn treatment had the least wasp emergence of the three treatments, while the total number of galls was not affected by treatment. Wasp mortality, not gall destruction, is why wasp emergence is reduced in burn treatments. More galls were located and, for the burn treatments, higher wasp emergence was seen from galls found within the leaf litter than those above it. Five co-inhabitants emerged from blueberry stem galls in this study. Three, Eurytoma solenozopheriae (Ashmead), Sycophila vacciniicola (Balduf), and Orymus vacciniicola (Ashmead) are commonly found associates. The other two, Eupelmus vesicularis (Ritzius) and Pteromalus spp., are new records for Nova Scotia. O. vacciniicola is likely an inquiline because it is the largest wasp emerging from galls, and there was a positive relationship between its emergence and that of H. nubilipennis. Larger gall size improved H. nubilipennis emergence from mow and spring-burn galls. After a field has been mowed in the fall, we recommend a spring burn to reduce gall populations and the threat of product contamination.     

Effect of gall formation by a cynipid wasp, Andricus symbioticus, on the development of the leaves and shoots of Quercus dentata

The sexual generation of a cynipid wasp, Andricus symbioticus Kovalev, forms its leaf galls most frequently near and on the leaf petiole of Quercus trees. I examined the effect of gall formation by A. symbioticus on the leaf development of a host plant, Quercus dentata Thunberg, by comparing the size and shape of galled and ungalled leaves. I also examined the effect of gall formation on shoot development by comparing the length of shoots with and without galled leaves. Three of seven Q. dentata trees surveyed were heavily infested with A. symbioticus. Leaf size did not differ between galled and ungalled leaves. However, the ratio of leaf width to length was greater in galled leaves, which is regarded to be a result of gall formation by A. symbioticus inhibiting the growth in length of Q. dentata leaves. Shoot length did not differ significantly between shoots with and without galled leaves. These results suggest that galls of A. symbioticus act as a sink that competes with leaves for reserved photoassimilates.     

Non-random distribution among a guild of parasitoids: implications for community structure and host survival

Abstract 1. Immature stages of the gall midge, Asphondylia borrichiae, are attacked by four species of parasitoids, which vary in size and relative abundance within patches of the gall midge’s primary host plant, sea oxeye daisy (Borrichia frutescens). 2. In the current study, a bagging experiment found that the smallest wasp, Galeopsomyia haemon, was most abundant in galls exposed to natural enemies early in the experiment, when gall diameter is smallest, while the wasp with the longest ovipositor, Torymus umbilicatus, dominated the parasitoid community in galls that were not exposed until the 5th and 6th weeks when gall diameter is maximal. 3. Moreover, the mean number of parasitoids captured using large artificial galls were 70% and 150% higher compared with medium and small galls respectively, while stem height of artificial galls significantly affected parasitoid distribution. Galls that were level with the top of the sea oxeye canopy captured 60% more parasitoids compared with those below the canopy and 50% more than galls higher than the plant canopy. 4. These non‐random patterns were driven primarily by the differential distribution of the largest parasitoid, T. umbilicatus, which was found significantly more often than expected on large galls and the smallest parasitoid of the guild, G. haemon, which tended to be more common on stems level with the top of the plant canopy. 5. Large Asphondylia galls, especially those located near the top of the Borrichia canopy, were more likely to be discovered by searching parasitoids. Results using artificial galls were consistent with rates of parasitism of Asphondylia galls in native patches of sea oxeye daisy. Gall diameter was 19% greater and the rate of parasitism was reduced by almost 50% on short stems; as a result, gall abundance was 24% higher on short stems compared with ones located near the top of the plant canopy. 6. These results suggest that parasitoid community composition within galls is regulated by both interspecific differences in ovipositor length and preferences for specific gall size and/or stem length classes.     

Abundance of Gall Midges on Poulsenia armata (Moraceae): Importance of Host Plant Size and Light Environment in Tropical Rain Forests1

We studied the effects of contrasting light environments on the relationship between the host plant size of Poulsenia armata and the abundance of two gall midges in a tropical rain forest in Veracruz, Mexico. The number and density of two gall morphs (i.e., laminar and vein‐petiole galls) were positively correlated with plant size only in trees found in the forest but not in gaps. The availability of foliar area of P. armata trees was greater in forest gaps than in the forest. The foliar area was positively correlated with the abundance of laminar galls in trees in forest sites, but not with vein‐petiole galls. We concluded that the abundance of two morphs of gall midges on P. armata was associated with host plant size only in the forest trees. Larger plants had more galls than small plants, although this relationship was affected by local light environments.     

Impact of whole-canopy and systemic insecticidal treatments on Callirhytis cornigera (Hymenoptera: Cynipidae) and associated parasitoids on pin oak

The gall wasp Callirhytis cornigera (Osten Sacken) is a cynipid with alternating generations that produce large, woody stem galls and tiny blister-like leaf galls on pin oak, Quercus palustris Muenchhausen, in the United States. We tested 3 approaches to control the leaf-galling generation, and determined their impact on associated parasitoids and effectiveness in reducing numbers of new stem galls. First, trees were sprayed with bifenthrin or chlorpyrifos in late March to kill females emerging from stem galls before they oviposited into buds. Second, concentrated solutions of abamectin, imidacloprid, or bidrin were injected from pressurized containers into tree sapwood to control larvae developing in young leaf galls. Finally, systemic insecticides (acephate, abamectin, dimethoate, or imidacloprid) were sprayed at early leaf expansion (2 May) or to young, expanded leaves (17 May) to target larvae in leaf galls. Parasitoids, mostly eulophids, accounted for approximately 70% mortality of leaf-galling C. cornigera larvae on untreated trees. Whole-canopy sprays during C. cornigera emergence from stem galls reduced overall numbers of galled leaves and leaf galls. Trunk injections of bidrin or abamectin resulted in significant mortality of gall inhabitants, including parasitoids. However, neither of the aforementioned approaches significantly reduced numbers of new stem galls. Sprays of abamectin, dimethoate, or imidacloprid applied on 2 May caused high mortality of all gall inhabitants. There was no net benefit, however, because parasitism caused a similar reduction in C. cornigera survival on unsprayed shoots. Sprays applied later in leaf expansion had little impact on gall inhabitants. Of the treatments tested, bifenthrin sprays at bud break provided the greatest reduction in new leaf galls, whereas bidrin injections provided the greatest reduction in gall wasps emerging from galled leaves. This study suggests that gall wasp outbreaks are unlikely to be controlled by a single treatment, regardless of application method.     

瘿绵蚜科虫瘿的多样性研究

是否形成虫瘿及虫瘿的位置、形态等是蚜虫生物学的重要特征.本文在已有标本采集记录和资料的基础上,从形成虫瘿的寄主植物、虫瘿的类型、虫瘿着生的部位和虫瘿的形态结构4方面对瘿绵蚜的虫瘿多样性进行了系统研究.结果表明该科蚜虫的虫瘿在类型上有虫瘿和伪虫瘿之别;在着生部位上,有叶片、侧脉、主脉、叶柄、复叶总轴、嫩枝等不同部位;就虫瘿而言,在形状上有尖椒状、鸡冠状、袋状、球状、黄瓜状、枫叶状和倍花状之分,在结构上有单室和多室两类.同时,基于系统分类及虫瘿多样性研究的结果,初步探讨了瘿绵蚜科虫瘿的演化,为进一步从分子水平深入研究虫瘿演化奠定基础.     

Anatomical profiles validate gall morphospecies under similar morphotypes

Plant galls are generated by the stimuli of gall-inducing organisms on their hosts, creating gall morphotypes that vary in color, shape, size, and tissue organization. Herein, we propose to compare the structural features of gall morphotypes on the superhost Croton floribundus (Euphorbiaceae) in order to recognize gall morphospecies, i.e., galls with similar shapes but different internal structures. Non-galled leaves and galls were analyzed macroscopically, histologically, and histochemically for the detection of primary metabolites, and the results obtained were used for statistical analyses of similarity. Among the eight gall morphospecies, four are globoid, two are lenticular, one is fusiform and one is marginal leaf rolling. Stomatal differentiation and the occurrence of different types of trichomes were impaired in some gall morphospecies. Three patterns of organization of the ground system are recognized, ranging from the maintenance of mesophyll cells that differentiate into palisade and spongy cells dorsiventrally to the formation of a complex cortex with three morphofunctional layers. The marginal leaf rolling galls have the simplest anatomical structures, quite similar to those of the non-galled host leaf, while lenticular, globoid (types I to IV), and fusiform galls are anatomically more complex. Herein, we report on eight gall morphospecies occurring on C. floribundus, which are distinguished by morpho-anatomical attributes and show the disruption of the morphogenetic patterns of the host leaf toward the morphogenesis of unique gall features.

     

The chemical composition of plant galls: are levels of nutrients and secondary compounds controlled by the gall-former?

The chemical composition of galled and ungalled plant tissue was compared in a series of experiments. Gall and adjacent plant tissue was analysed for 20 species of gall-former on 11 different plant species. There were clear differences between galled and ungalled tissue in levels of nutrients and secondary compounds. Gall tissue generally contained lower levels of nitrogen and higher levels of phenolic compounds than ungalled plant tissue. The gall tissue produced by the same plant in response to different species of gall-former differed in chemical composition, as did the gall-tissue from young and mature galls of the same species. The chemical differences between gall and plant tissues were studied in more detail in two field manipulations. Firstly, the seasonal changes in phenolic biosynthesis in Pontania proxima and P. pedunculi (Hymenoptera: Tenthredinidae) gall tissue were compared to those of their host plants, Salix alba and S. caprea. In both types of gall tissue, phenolic levels declined as the season progressed, but levels in the surrounding plant tissue increased. When the gall insects were killed with insecticide, phenolic levels in the galled tissue dropped to the same level as those in adjacent plant tissue. Secondly, the density of Cynips divisa (Hymenoptera: Cynipidae) galls on Quercus robur leaves was reduced by removing half the galls present, either those from the central region of the leaf or those from the edge. Decreasing gall density increased the size of the remaining galls and the weight of the insects, but these effects were most marked when the galls remaining were growing centrally on the leaf, i.e. when the galls from the edge had been removed. Decreasing gall density increased the nitrogen content of the remaining galls, again to a greater extent in galls growing centrally on the leaf. The results of these studies suggest that the levels of nutrients and secondary compounds in gall tissue are usually markedly different to those of surrounding plant tissue, and that gall-formers may produce species-specific and temporally variable changes in the chemical composition of gall tissue. Received: 7 July 1997 / Accepted: 29 September 1997     

Preference–performance relationship in the gall midge Rabdophaga rosaria: insights from a common‐garden experiment with nine willow clones

1. Oviposition preferences of herbivorous insects are predicted to match offspring performance on different host taxa or on conspecific host genotypes. In gall‐inducing insects, host‐plant properties such as growth rate and gall size, which are determined by plant genotype and growing conditions, may have a significant impact on offspring performance and, hence, should influence oviposition site selection. 2. The present study investigated host preference of the European rosette willow gall midge Rabdophaga rosaria (Loew) in relation to offspring success on seven clones of Salix myrsinifolia Salisb. and two naturally hybridised S. myrsinifolia × phylicifolia L. clones growing in a replicated design in an experimental field under two fertilisation regimes. For each clone, the average growth rate, number of shoot tips, and leaf and gall size were determined, and their effects on midge preference and larval survival were examined. 3. Main shoot height, number of shoot tips, and gall size were significantly affected by clone. The midges clearly preferred certain clones over the others, but preferences were not related to willow growth traits or to gall size. Survival probability was higher in large than in small galls, but females did not prefer large‐leaved clones that produced the biggest rosette galls. Midge oviposition was also uncorrelated with prior rates of leaf‐rust infection and with feeding preferences of voles and folivorous insects. 4. The weak preference–performance relationship of R. rosaria within S. myrsinifolia is probably explained by evolutionary constraints that prevent generalist insects from achieving an ability to discriminate among conspecific hosts of variable quality.     

Oxycephalomyia, gen. nov., and life history strategy of O. styraci comb. nov. (Diptera: Cecidomyiidae) on Styrax japonicus (Styracaceae)

A new genus Oxycephalomyia is described to contain the gall midge that was previously known as Asteralobia styraci (Shinji). Oxycephalomyia styraci, comb. nov., produces leaf vein galls on Styrax japonicus (Styracaceae). The adult of O. styraci is redescribed, and its full‐grown larva and pupa are described for the first time. The annual life cycle of the gall midge in northern Kyushu was clarified; the first instars overwinter in the galls on the host plant. However, the galls of O. styraci mature much later in the season than those of other gall midges with a similar life history pattern, and the durations of second and third larval instars are remarkably short. Such a life history pattern is considered to have an adaptive significance in avoiding larval parasitism, particularly by early attackers. The number of host axillary buds as oviposition sites decreased in bearing years and increased in off years, but there was no sign of oviposition site shortage even in bearing years, probably due to the low population density of the gall midge. An unidentified lepidopteran that feeds on galled and ungalled host buds and a Torymus sp. that attacks pupae of O. styraci were recognized as mortality factors of the gall midge.     

Invasion by the chestnut gall wasp in Italy causes significant yield loss in Castanea sativa nut production

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Tritrophic interactions and trade-offs in herbivore fecundity on hybridising host plants

Abstract. 1. Interspecific plant hybridisation can have important evolutionary consequences for hybridising plants and for the organisms that they interact with on multiple trophic levels. In this study the effects of plant hybridisation on the abundance of herbivores and on the levels of herbivore parasitism were investigated. 2. Borrichia frutescens, B. arborescens, and their hybrid (B. × cubana) were censused for Asphondylia borrichiae galls and Pissonotus quadripustulatus plant hoppers in the Florida Keys. Levels of egg parasitism were determined by dissecting parental and hybrid stems and galls for herbivore and parasite eggs and larvae. Stem toughness and gall size are plant‐mediated modes of protection from parasitism and these were also measured. For gall midges, fly size was measured as an estimate of fecundity. 3. Field censuses indicated that herbivore abundances varied on hybrid hosts relative to parent plant species and that the different herbivore species exhibited different patterns of abundance. Asphondylia borrichiae gall numbers followed the additive pattern of abundance while P. quadripustulatus numbers most closely resembled the dominance pattern. 4. Parasitism of P. quadripustulatus eggs was high on B. frutescens and the hybrids, and low on B. arborescens, which also had significantly tougher stems. Asphondylia borrichiae suffered the highest levels of parasitism on B. frutescens, the host plant which produced the smallest galls. On B. arborescens, which produced the largest galls, levels of A. borrichiae parasitism were lowest. Both parasitism and gall size were intermediate on the hybrid plants. Galls on B. arborescens and hybrid plants produced significantly smaller flies then those from B. frutescens suggesting that, when selecting hosts from among parent species and hybrids, gall flies may face a trade‐off between escape from natural enemies and maximising fecundity.     

ANATOMY OF SOME LEAF GALLS OF ROSA WOODSII (ROSACEAE)

Infection of Rosa woodsii by some members of the order Hymenoptera results in neoplasmic outgrowths on the leaves. One type of outgrowth produces a spherical swelling (leaf gall) while the other has extensive hair-like proliferations (hairy gall). The anatomy and ultrastructure of these galls were examined by light microscopy and transmission electron microscopy. The leaf gall cells were considerably larger than normal cells, lacked well-developed chloroplasts and were loosely arranged with prominent intercellular spaces. Vascular bundles were scattered throughout the gall tissue. The upper three cell layers of the leaf gall tissue resembles a periderm, having many suberin lamellae. The suberin lamellae were often traversed by pores which may represent incomplete plasmodesmata. Phenolic compounds were commonly seen both in the normal and gall cells. A layer of internal cells of the hairy galls have remarkably thickened cell walls, presumably due to the deposition of cellulosic substances. Unlike leaf galls, the epidermal cells of the hairy galls were not heavily cuticularized and no periderm was found. The hair-like outgrowths present on the outer surface of these galls had a central vascular bundle. The epidermis of the outgrowths also had thickened cell walls, and trichomes occurred on the outer surface. The structural modifications brought about by the insect invasion in these two galls are compared and their roles in gall formation are discussed.