Phenotypic plasticity and similarity among gall morphotypes on a superhost,Baccharis reticularia (Asteraceae)

Understanding factors that modulate plant development is still a challenging task in plant biology. Although research has highlighted the role of abiotic and biotic factors in determining final plant structure, we know little of how these factors combine to produce specific developmental patterns. Here, we studied patterns of cell and tissue organisation in galled and non‐galled organs of Baccharis reticularia, a Neotropical shrub that hosts over ten species of galling insects. We employed qualitative and quantitative approaches to understand patterns of growth and differentiation in its four most abundant gall morphotypes. We compared two leaf galls induced by sap‐sucking Hemiptera and stem galls induced by a Lepidopteran and a Dipteran, Cecidomyiidae. The hypotheses tested were: (i) the more complex the galls, the more distinct they are from their non‐galled host; (ii) galls induced on less plastic host organs, e.g. stems, develop under more morphogenetic constraints and, therefore, should be more similar among themselves than galls induced on more plastic organs. We also evaluated the plant sex preference of gall‐inducing insects for oviposition. Simple galls were qualitative and quantitatively more similar to non‐galled organs than complex galls, thereby supporting the first hypothesis. Unexpectedly, stem galls had more similarities between them than to their host organ, hence only partially supporting the second hypothesis. Similarity among stem galls may be caused by the restrictive pattern of host stems. The opposite trend was observed for host leaves, which generate either similar or distinct gall morphotypes due to their higher phenotypic plasticity. The Relative Distance of Plasticity Index for non‐galled stems and stem galls ranged from 0.02 to 0.42. Our results strongly suggest that both tissue plasticity and gall inducer identity interact to determine plant developmental patterns, and therefore, final gall structure.     

Linking Avicennia germinans (Acanthaceae) architecture to gall richness and abundance in Brazilian Amazon mangroves

The diversity and abundance of gall‐inducing organisms are directly proportional to the structural complexity of the host plant. This hypothesis is controversial for forest environments, such as mangroves. Avicennia germinans (L.), a principal mangrove tree species found in the Neotropical region, is considered to be a superhost for gall‐inducing insects. Using a generalized linear mixed model (GLMM) based on the analysis of 1000 apical branches from 50 A. germinans trees, we examined the diversity and abundance of gall morphotypes (GM), together with the structural attributes of replanted 5‐ to 9‐year‐old mangroves, in the Amazon coast of Brazil. A total of 7602 galls were registered, averaging 1.3 ± 0.4 galls per leaf. Sixteen of the 22 morphotypes identified were found at all study sites. Two gall morphotypes (GM7 and GM4) were the most abundant, representing approximately 40 percent of the total. The structural complexity of the plant (mainly based on the number of leaves) directly affected the abundance and diversity of these organisms. While A. germinans is a superhost, this type of parasitism did not affect plant development or survival. The ample distribution of A. germinans, the formation of monospecific forests, and the high palatability of this plant make it an essential resource for the survival of the gall‐inducing guild in the mangroves of the Neotropics.     

Partitioning of herbivore hosts across time and food plants promotes diversification in the Megastigmus dorsalis oak gall parasitoid complex

Communities of insect herbivores and their natural enemies are rich and ecologically crucial components of terrestrial biodiversity. Understanding the processes that promote their origin and maintenance is thus of considerable interest. One major proposed mechanism is ecological speciation through host‐associated differentiation (HAD), the divergence of a polyphagous species first into ecological host races and eventually into more specialized daughter species. The rich chalcid parasitoid communities attacking cynipid oak gall wasp hosts are structured by multiple host traits, including food plant taxon, host gall phenology, and gall structure. Here, we ask whether the same traits structure genetic diversity within supposedly generalist parasitoid morphospecies. We use mitochondrial DNA sequences and microsatellite genotypes to quantify HAD for Megastigmus (Bootanomyia) dorsalis, a complex of two apparently generalist cryptic parasitoid species attacking oak galls. Ancient Balkan refugial populations showed phenological separation between the cryptic species, one primarily attacking spring galls, and the other mainly attacking autumn galls. The spring species also contained host races specializing on galls developing on different host‐plant lineages (sections Cerris vs. Quercus) within the oak genus Quercus. These results indicate more significant host‐associated structuring within oak gall parasitoid communities than previously thought and support ecological theory predicting the evolution of specialist lineages within generalist parasitoids. In contrast, UK populations of the autumn cryptic species associated with both native and recently invading oak gall wasps showed no evidence of population differentiation, implying rapid recruitment of native parasitoid populations onto invading hosts, and hence potential for natural biological control. This is of significance given recent rapid range expansion of the economically damaging chestnut gall wasp, Dryocosmus kuriphilus, in Europe.     

The role of pectic composition of cell walls in the determination of the new shape-functional design in galls of Baccharis reticularia (Asteraceae)

The pectic composition of cell wall is altered during the processes of cell differentiation, plant growth, and development. These alterations may be time-dependent, and fluctuate in distinct regions of the same cell or tissue layer, due to the biotic stress caused by the activity of the gall inducer. Among the roles of the pectins in cell wall, elasticity, rigidity, porosity, and control of cell death may be crucial during gall development. Galls on Baccharis reticularia present species-specific patterns of development leading to related morphotypes where pectins were widely detected by Ruthenium red, and the pectic epitopes were labeled with specific monoclonal antibodies (LM1, LM2, LM5, LM6, JIM5, and JIM7) in distinct sites of the non-galled and the galled tissues. In the studied system B. reticularia, the epitopes for extensins were not labeled in the non-galled tissues, as well as in those of the rolling and kidney-shaped galls. The high methyl-esterified homogalacturonans (HGA) were labeled all over the tissues either of non-galled leaves or of the three gall morphotypes, while the intense labeling for arabinogalactans was obtained just in the rolling galls. The pectic composition of non-galled leaves denotes their maturity. The kidney-shaped gall was the most similar to the non-galled leaves. The pectic dynamics in the gall tissues was particularly altered in relation to low methyl-esterified HGA, which confers elasticity and expansion, as well as porosity and adhesion to cell walls, and are related to the homogenization and hypertrophy of gall cortex, and to translocation of solutes to the larval chamber. Herein, the importance of the pectic dynamics of cell walls to the new functional design established during gall development is discussed for the first time. The repetitive developmental patterns in galls are elegant models for studies on cell differentiation.     

Histocytological aspects of four types of ambrosia galls on Machilus zuihoensis Hayata (Lauraceae)

Four types of prosoplasmatic galls induced by Daphnephila midges are found on leaves of Machilus zuihoensis, a species endemic to Taiwan: urn- and small urn-shaped, obovate, and hairy oblong galls. In addition to containing nutritive tissues, these galls are lined with fungal hyphae. The objective of this study was to describe and compare the structural organization of the various gall morphologies and to examine the ultrastructure of the nutritive and fungal cells lining the gall chambers. The morphology and ultrastructure of mature-stage galls were examined by light, scanning electron, and transmission electron microscopy. Diverse epidermal cell shapes and wax textures were observed in the leaves and galls of M. zuihoensis. In small urn-shaped, obovate, and hairy oblong galls vascular bundles extend from the gall base to near the centre of the gall top. In contrast, vascular bundles in urn-shaped galls are distributed in the gall wall and extend to close to the outer gall top. Trichomes were present only abaxially on leaves and on hairy oblong gall surfaces. Starch granules, tannins, and mucilage were distributed differently among the four gall types. Further, fungal mycelia spread in the interior gall wall and partially passed through the intercellular spaces of nutritive cells and reached the sclerenchyma. Histological analyses revealed that the surface structure of galls differs from that of the leaf and that the epidermal organization differs among the four gall types. Different types of leaf galls on the same plant have different patterns of tissue stratification and contain different ergastic substances. The results of this study will contribute to the understanding of tritrophic relationships and the complex interactions among parasitic gall-inducing insects, mutualistic fungi, and host plants.     

Developmental anatomy and immunocytochemistry reveal the neo-ontogenesis of the leaf tissues of Psidium myrtoides (Myrtaceae) towards the globoid galls of Nothotrioza myrtoidis (Triozidae)

Key message

The temporal balance between hyperplasia and hypertrophy, and the new functions of different cell lineages led to cell transformations in a centrifugal gradient that determines the gall globoid shape.

Abstract

Plant galls develop by the redifferentiation of new cell types originated from those of the host plants, with new functional and structural designs related to the composition of cell walls and cell contents. Variations in cell wall composition have just started to be explored with the perspective of gall development, and are herein related to the histochemical gradients previously detected on Psidium myrtoides galls. Young and mature leaves of P. myrtoides and galls of Nothotrioza myrtoidis at different developmental stages were analysed using anatomical, cytometrical and immunocytochemical approaches. The gall parenchyma presents transformations in the size and shape of the cells in distinct tissue layers, and variations of pectin and protein domains in cell walls. The temporal balance between tissue hyperplasia and cell hypertrophy, and the new functions of different cell lineages led to cell transformations in a centrifugal gradient, which determines the globoid shape of the gall. The distribution of cell wall epitopes affected cell wall flexibility and rigidity, towards gall maturation. By senescence, it provided functional stability for the outer cortical parenchyma. The detection of the demethylesterified homogalacturonans (HGAs) denoted the activity of the pectin methylesterases (PMEs) during the senescent phase, and was a novel time-based detection linked to the increased rigidity of the cell walls, and to the gall opening. Current investigation firstly reports the influence of immunocytochemistry of plant cell walls over the development of leaf tissues, determining their neo-ontogenesis towards a new phenotype, i.e., the globoid gall morphotype.     

Arthropod-induced neoplastic formations on trees change photosynthetic pigment levels and oxidative enzyme activities

Abstract

In order to test the hypothesis that arthropod-induced neoplastic formations on trees affect biochemical characteristics of both the newly formed galls and host plant tissues, biochemical characteristics with a possible adaptive role were determined in nine gall-former–host tree combinations. Photosynthetic pigments, extractable protein content, and oxidative enzyme activities were determined in gall tissues, leaf tissues of galled leaves, and leaves on ungalled tree branches. Neoplastic tissues were characterized by a low content of photosynthetic pigments, decreased chlorophyll a/b ratio, lower extractable protein content, and decreased activities of peroxidase and polyphenol oxidase as compared with ungalled host leaf tissues. In galled leaves or in leaves adjacent to galls, increased level of peroxidase activity was found. In several gall-inducer–host plant combinations, galled host plant tissues contained increased activity of polyphenol oxidase as well. The presented data reflect long-term systemic effects of neoplastic formation on host tree physiology suggesting that gall inducers affect potential adaptive responses of host plants.     

Manipulation of food resources by a gall-forming aphid: the physiology of sink-source interactions

Summary We examined the capacity of the galling aphid, Pemphigus betae, to manipulate the sink-source translocation patterns of its host, narrowleaf cottonwood (Populus angustifolia). A series of ¹⁴C-labeling experiments and a biomass allocation experiment showed that P. betae galls functioned as physiologic sinks, drawing in resources from surrounding plant sources. Early gall development was dependent on aphid sinks increasing allocation from storage reserves of the stem, and later development of the progeny within the gall was dependent on resources from the galled leaf blade and from neighboring leaves. Regardless of gall position within a leaf, aphids intercepted ¹⁴C exported from the galled leaf (a non-mobilized source). However, only aphid galls at the most basal site of the leaf were strong sinks for ¹⁴C fixed in neighboring leaves (a mobilized source). Drawing resources from neighboring leaves represents active herbivore manipulation of normal host transport patterns. Neighboring leaves supplied 29% of the ¹⁴C accumulating in aphids in basal galls, while only supplying 7% to aphids in distal galls. This additional resource available to aphids in basal galls can account for the 65% increase in progeny produced in basal galls compared to galls located more distally on the leaf and limited to the galled leaf as a food resource. Developing furits also act as skins and compete with aphid-induced sinks for food supply. Aphid success in producing galls was increased 31% when surrounding female catkins were removed.     

Floral-like destiny induced by a galling Cecidomyiidae on the axillary buds of Marcetia taxifolia (Melastomataceae)

The galls induced by Cecidomyiidae, Diptera, are very diverse, with conspicuous evidence of tissue manipulation by the galling herbivores. Bud galls, as those induced by an unidentified Cecidomyiidae species on Marcetia taxifolia, Melastomataceae, can be considered as one of the most complex type of prosoplasma galls. The gall-inducer manipulate the axillary meristem of the plant in a way that gall morphogenesis may present both vegetative and reproductive features of the host plant. Herein, we analyzed traces of determinate and indeterminate growth in the bud gall of M. taxifolia, looking for parallels between the features of the leaves and flowers, natural fates of the meristematic cells. The bud galls are induced by the cecidomyiid fly, and are formed by the connation of eight leaf primordia, a common process in ovary morphogenesis. The bud gall corresponds to a pistil-shaped gall morphotype, with anatomical features similar to those of an hypanthium and sepals. The gall mimics an ovary, which has protective barriers at the apex, and a nutritive tissue (with storage of lipids and proteins) or a placenta, respectively, at the basal portion. The redifferentiation of the promeristem into a nutritive tissue at the base of the gall confers a determinate destiny to the axillary bud. Comparatively, the gradients of cell expansion and of accumulation of primary metabolites also indicate that the gall and the ovary are convergent structures. Some constraints of the host plant cells, such as the absence of lignification, and the accumulation of polyphenols, lipids and terpenoids, are not altered and may confer chemical protection for plant tissues and the larva against oxidative stress.     

Carnivores and carotenoids are associated with adaptive behavioural divergence in a radiation of gall midges

1. Adaptive divergence in sympatry is supposed to be inhibited by the homogenizing role of gene flow. However, studies continue to uncover examples of sympatric divergence. In this study, two divergent phenotypes in a complex of four syntopic gall midge morphotypes [nominally Asteromyia carbonifera Osten Saken, Diptera: Cecidomyiidae: Alycaulini] are characterised. The first is a behavioural phenotype governing within‐host tissue preference and the second is a trait governing accessory‐gland carotenoid quality and quantity. 2. One gall morphotype (crescents) lay most of their eggs on mature tissue whereas the other three gall morphotypes oviposit only on young emerging leaves. Ecological maintenance of this divergent trait appears to be driven by enemy‐reduced space. That is, nearly 40% of the crescent morphotype galls that develop high on the plant are attacked by the egg parasitoid Platygaster solidaginis Ashmed, whereas those low on the plant are relatively protected. 3. All morphotypes contain carotenoids in their accessory glands, but the quality and quantity of these pigments differs significantly between the morphotypes and is positively associated with the probability of parasitism by P. solidaginis. 4. Larval salivary glands also contain carotenoids and the plant hormone abscisic acid, which in plants is synthesized from carotenoid precursors and is involved in regulating plant defences. This hormone may facilitate gall development and influence gall morphology. 5. Ecological fitness trade‐offs between carotenoids, parasitoid attack, and plant resistance may be fostering adaptive divergence in ovipositional phenotypes and sympatric speciation in this complex of gall midge morphotypes.     

Red galls: the different stories of two gall types on the same host

• Several studies have suggested reasons why galls have conspicuous colours, but none of the ideas have been confirmed. However, what if the vibrant colours of some galls are explained simply by the effect of light exposure? This may lead to anthocyanin accumulation, functioning as a defence mechanism against the effects of high light.
• We studied the globoid galls induced by Cecidomyiidae (Diptera) on Qualea parviflora (Vochysiaceae), relating anthocyanin accumulation and chlorophyll fluorescence parameters to light incidence in abaxial and adaxial galls. We also tested if the anthocyanin accumulation patterns apply to another Cecidomyiidae‐induced gall morphotype (intralaminar) within the same plant.
• Adaxial galls are exposed to higher incident light, with more anthocyanin accumulation and therefore red coloration. In galls from angled leaves, the greater the angle of the leaf, the higher the difference between anthocyanins on the sun and shade sides of galls. Photosynthetic pigment concentrations did not differ between abaxial and adaxial galls. However, we found higher (F_m′ ? F′)/F_m′ and F_v/F_m in the abaxial galls. Conversely, NPQ and R_fd were higher in adaxial galls. Finally, the pattern of anthocyanin accumulation was not found in the intralaminar gall.
• Anthocyanin accumulation in galls functions as a photoprotective strategy, maintaining tissue vitality in regions exposed to high light conditions. However, this mechanism may vary even among galls within the same host, indicating idiosyncrasy when it comes to coloration in galls. To date, this is the first study to demonstrate quantitatively why the galls of a specific species may be coloured: the variation in light regimes creates differential anthocyanin accumulation, influencing coloration.

     

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     

Organization of a parasitoid community associated with a complex of galls on Atriplex spp. in southern California

Abstract. 1. Atriplex canescens (Pursh) Nuttall and A.polycarpa (Torrey) Watson (Chenopodiaceae) support twelve morphologically distinct gall types in southern California. Thirty-seven common species of parasitoids, predators and inquilines are associated with these galls. 2. The galls incited by eight members of the Asphondylia atriplicis Cockerell (Diptera: Cecidomyiidae) species complex are linked into a single, interacting community through shared hymenopterous parasitoids and inquilines. 3. Cluster analysis (UPGMA) grouped the fifteen most common species of Chalcidoidea into three host guilds of five species each: (1) specialists in tumour stem and blister leaf galls on A.canescens, (2) specialists in woolly stem galls on A.poiycarpa, and (3) generalists that attack all galls. Guild 1 dominated the galls with which it was primarily associated, while guild 3 dominated the remainder. 4. The abundances of the parasitoids of the tumour stem and blister leaf galls were negatively correlated with the abundances of two organizer species, a gall-forming inquiline, Tetrastichus cecidobroter Gordh and Hawkins, and an internal, larval—pupal parasitoid, Tetrastichus sp. B. The abundances of nine of the twelve most common chalcidoids were not correlated with the abundances of all coaccurring species in six other galls. 5. Host seasonality partly determines parasitoid population dynamics and guild structure. Parasitoid dominance increased with gall duration, suggesting that parasitoid competition depends on resource stability. The two continuously available galls were dominated by their specialist guild, while all seasonal galls were dominated by generalists. The subdominant specialists of woolly stem galls may represent competitively inferior species that utilize those galls opportunistically, because of the gall's widespread distribution and 9–10 month yearly availability. 6. Sites in the Colorado Desert and chaparral that supported several gall types showed stable relative abundances of the major parasitoid species, whereas sites in the Mojave Desert that supported only woolly stem galls had unpredictable parasitoid species assemblages. 7. The competitive success of Atriplex gall parasitoids may depend primarily on voltinism (multivoltine species dominated univoltine species) and mode of feeding (phytophagous, mixed entomophagous—phytophagous and facultatively hyperparasitic species in general dominated strict primary parasitoids).     

Cytological and histochemical gradients on two <Emphasis Type="BoldItalic">Copaifera langsdorffii</Emphasis> Desf. (Fabaceae)—Cecidomyiidae gall systems

Previous ultrastructural and histochemical analysis proposed patterns in the accumulation of substances in galls of Diptera: Cecidomyiidae in some plant species of the temperate region. Similar analyses were done to verify the conservativeness of these patterns in the Neotropical region, where a great number of species of Cecidomyiidae is responsible for a wide diversity of morphotypes. Two gall morphotypes induced by Cecidomyiidae in a unique host plant, Copaifera langsdorffii, were studied. The gradients of carbohydrates and the activity of invertases and acid phosphatases were similar, but the cytological gradients and distribution of proteins evidenced that the sites of the induction as well as the amount of neoformed tissues may be peculiar to each gall system. The production of lipids just in the secretory cavities either in the non-galled or galled tissues indicated a potentiality of the host plant which could not be manipulated by the galling insects. Further, the absence of nucleus in the nutritive tissue, an exclusive feature of the horn-shaped galls, indicates cell death attributed to the feeding habit of the galling herbivore.     

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.     

Gall distribution as a compromise between the optimal gall-site selection and the synchrony to host-plant phenology in the aphid Kaltenbachiella japonica

Gall-site selection by the aphid Kaltenbachiella japonica was evaluated in relation to leaf position in a shoot, and gall positions within a leaf. First-instar fundatrices induce closed galls on the midribs of host leaves, and several galls were often induced on one leaf. Leaves with many galls were often withered before emergence of sexuparae from the galls. Within a leaf, gall volume was positively correlated with the sum of lateral-vein length in the leaf segment at which the gall was induced. The observed pattern in gall volume among the leaf segments corresponded with that in the lateral-vein length. These results show that a foundatrix selects the most vigorous position within a leaf to produce more offspring. Although distal leaves grew faster than did basal leaves, gall density was highest on leaves at the middle order when a shoot has more than seven leaves. Optimal gall-site selection seems to be constrained by the asynchrony in timing between the hatching of fundatrices and leaf growth within a shoot. These results suggest that the observed gall distribution is affected by both the distribution of suitable galling sites within a leaf and the synchrony with leaf phenology of the host plant.     

Leaf‐derived cecidomyiid galls are sinks in Machilus thunbergii (Lauraceae) leaves

Three relevant hypotheses – nutrition, environment and the enemies hypothesis – often invoked to explore source and sink relationships between galls and their host plants are still under dispute. In this research, chlorophyll fluorescence, gas exchange capacity, stomatal conductance, total carbon and nitrogen, total soluble sugars and starches, and scanning and transmission electron microscopy of two types of galls were used to investigate source–sink relationships. Compared with host leaves, these galls demonstrated slightly lower chlorophyll fluorescence; however, gas exchange capacity and stomatal conductance were not detected at all. Scanning electron micrographs demonstrated that the abaxial epidermis of host leaves contain normal amounts of stomata, whereas no stomata were observed on the exterior and interior surfaces of both types of galls. In addition, gall inner surfaces were covered with many kinds of fungal hyphae. Gall total carbon (C) and nitrogen (N) levels were lower but the C/N ratio was higher in galls than host leaves. Both types of galls accumulated higher total soluble sugars and starches than host leaves. Transmission electron micrographs also revealed that both types of galls contain plastoglobuli and giant starch granules during gall development. Results strongly indicate that leaf‐derived cecidomyiid galls are sinks in Machilus thunbergii leaves. However, it is perplexing how larvae cycle and balance CO₂ and O₂ in gall growth chambers without stomata.     

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.     

Gall‐forming insects in a lowland tropical rainforest: low species diversity in an extremely specialised guild

1. Gall‐forming insects are a guild of endophages that exhibit a high level of fidelity to their host plants, however, their level of host specificity is seldom explicitly tested. 2. Gall‐forming insect taxa from 32 species of woody tropical plants with resolved phylogenetic relationships were collected and reared, representing 15 families from all the major clades of angiosperms, at three lowland rainforest locations in Madang, Papua New Guinea (PNG). 3. More than 8800 galled plant parts were collected from 78 gall morphospecies at an average of 2.4 per host plant. Total species richness at the sampling sites was estimated to be 83–89. All but one morphospecies were monophagous resulting in an effective specialisation of 0.98. 4. Specific leaf weight, foliar nitrogen, the presence of latex, and the successional preference of plant species all gave a phylogenetic signal, but only plant successional preference influenced the species richness of galls on analysis of phylogenetically independent contrasts. Gall species were distributed randomly among host plant species and showed no preference for any particular plant lineage. Furthermore, most gall‐forming taxa were evenly dispersed across the host plant phylogeny. 5. In the tropical rainforests of New Guinea, gall‐forming insects are ubiquitous but occur in species‐poor assemblages. Local species richness is closely tied to the diversity of angiosperms owing to very high host specificity. 6. Finally, galler species richness data from the literature across habitats and latitudes were compared and suggest that tropical rainforests may be richer in galls than previously acknowledged.     

Bottom‐up manipulations alter the community structures of galling insects and gall morphs on Neoboutonia macrocalyx trees in a moist tropical rainforest

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