In situ occurrence of a gall midge (Insecta,Diptera, Cecidomyiidae) on fossilized angiosperm leaf cuticle fragments from the Pliocene sediments of eastern India

In situ preservation of fossil insect damage in plant fossils is an excellent tool to study the coevolution of flora and fauna through geological time, but finding both damage and the insect causing that damage in the same specimen is a very rare phenomenon. Galling is a common form of angiosperm leaf damage, which can be regarded as a kind of extended phenotype of the causal insects, essentially the gall midges, but galls usually lack remains of the insects themselves. Here we report the in situ occurrence of a gall midge (Insecta, Diptera, Cecidomyiidae) as well as its pupal exuviae on the abaxial cuticular surface of fossilized leaf cuticle fragments of Fabaceae leaves (cf. Albizia) that also bear galls, recovered from the latest Neogene (Rajdanda Formation, Pliocene) sediments of the Chotonagpur Plateau, Jharkhand, northeastern India. This Pliocene gall midge features well-preserved legs, segmented antenna with distinct and enlarged scape, elongate curved setae, and longer than broad terminal plate of the ovipositor lamellae. The in situ presence of a gall midge on a host fabaceous leaf cuticle indicates the existence of a host-ectoparasite relationship in the ancient warm and humid tropical monsoon-influenced forests of eastern India during the Pliocene. This is the first authentic fossil record of an in situ phytophagous insect of Cecidomyiidae from India, as well as southeast Asia. Although the identification of the recovered phytophagous insect associated with the fossil leaf cuticle is only possible to family level, this find reveals that such plant-insect relationships existed in the Pliocene of eastern India.     

Pliocene Albizia (Fabaceae) from Jharkhand,eastern India: reappraisal of its biogeography during the Cenozoic in Southeast Asia

Albizia, a diverse tree genus, occupies monsoonal warm humid rain forests in tropical and subtropical regions. We recovered a well-preserved compound fossil leaf and two fossil fruits of Albizia (Fabaceae) from the latest Neogene (Rajdanda Formation: Pliocene) sediments of Jharkhand of the Chotanagpur Plateau, eastern India. On the basis of the architectural features of the fossil leaf, a new species is established as A. mahuadanrensis Hazra, Hazra and Khan, n. sp., characterised by a bipinnate, compound leaf having a rachis bearing opposite, asymmetrically ovate to sub-rhomboid leaflets, pulvinus on leaflet petiolule and brochiodromous secondary veins. Based on both morphological and anatomical characters of the fossil fruits, A. palaeoprocera Hazra, Hazra and Khan, n. sp. is erected, characterised by flattened to broadly linear shaped, wingless fruits; ovate-elliptic shaped seed chambers having ellipsoidal seeds in one series; irregularly polygonal to rectangular epidermal cells with oblique end walls and randomly oriented, scattered, paracytic stomata. Analysis of Albizia fossil occurrences indicates that the legume taxon was common in Neogene forests of India and elsewhere. The present-day distribution of the closely affiliated modern species of the fossil taxa indicates a warm and humid tropical environment during the time of deposition. We also review the biogeographic history of Albizia in India and other Asian countries.     

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.     

The number of cecidomyiid insect galls affects the photosynthesis of Machilus thunbergii host leaves

Previous studies of the impacts of galls on host leaf photosynthesis do not suggest any general trends, with a reported range of effects from negative to positive. In this study, photosynthetic characteristics such as chlorophyll fluorescence (Fv/Fm), photosynthetic capacity, and stomata conductance were determined in two types of fruit-like galls (red ovoid and green obovate galls) induced by Daphnephila taiwanensis and Daphnephila sueyenae, respectively, in order to investigate whether the number of galls affects the photosynthesis of galled leaves of Machilus thunbergii. In 2008, chlorophyll fluorescence and photosynthetic capacity were negatively correlated with gall numbers, non-significantly and significantly, respectively, whereas stomata conductance was positively but non-significantly correlated with gall numbers. In 2009, photosynthesis capacity and stomata conductance were negatively, but non-significantly, correlated with gall numbers. Results imply that photosynthesis in M. thunbergii leaves is slightly affected by the number of cecidomyiid insect galls, and that the higher the gall number, the greater the negative effect that galls have on host leaf photosynthesis and subsequent infection.     

Gall‐forming insect species richness along a non‐scleromorphic vegetation rainfall gradient in South Africa: The importance of plant community composition

Abstract Currently there is no single accepted hypothesis to explain gall‐forming insect species richness at a particular locality. Hygrothermal stress, soil nutrient availability, plant species richness, plant structural complexity, plant family or genus size, and host plant geographical range size have all been implicated in the determination of gall‐forming insect species richness. Previous studies of such richness at xeric sites have included predominantly scleromorphic vegetation, usually on nutrient‐poor soils. This study is the first to investigate gall‐forming insect species richness of xeric, non‐scleromorphic vegetation. Two habitat types were sampled at each of five localities across a rainfall gradient in the savanna biome of South Africa. The habitat types differed with respect to plant species composition and topography. Gall‐forming insect species richness did not increase with increasing hygrothermal stress or decreasing soil fertility. Rather, gall‐forming insect species richness was largely dependent on the presence of Terminalia sericea as well as other members of the Combretaceae and Mimosaceae. Plots where all these taxa were present had the highest gall‐forming insect species richness, up to 15 species, whereas plots with none of these taxa had a maximum of four galling‐insect species. Despite herb, shrub and tree strata not differing in gall‐forming insect species richness, insect galls were more common on woody than non‐woody plants. Also, stem galls were more frequent than apical or leaf galls. An alternative hypothesis to explain local gall‐forming insect species richness is suggested: galling insects may preferentially select those plant species with characteristics such as chemical toxicity, mechanical strength, degree of lignification or longevity that can be manipulated to benefit the galler. Thus plant community composition should be considered when attempting to explain gall‐forming insect species richness patterns.     

Fungal diversity in galls of baldcypress trees

The baldcypress midge (Taxodiomyia cupressi and Taxodiomyia cupressiananassa) forms a gall that originates from leaf tissue. Female insects may inoculate galls with fungi during oviposition, or endophytes from the leaf tissue may grow into the gall interior. We investigated fungal diversity inside of baldcypress galls, comparing the gall communities to leaves and comparing fungal communities in galls that had successful emergence versus no emergence of midges or parasitoids. Galls of midges that successfully emerged were associated with diverse gall fungal communities, some of which were the same as the fungi found in surrounding leaves. Galls with no insect emergence were characterized by relatively low fungal diversity.     

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.     

Host plant intraspecific variation determines gall traits

• Galls display a multiplicity of traits, including colours, which are driven by pigment accumulation. Their conspicuousness has attracted researchers' attention and several hypotheses have been raised. However, plants themselves vary intra-specifically, including in their pigment concentrations. As galls are a result of host tissue development, colours may be a by-product of the host's own traits, being more conspicuous simply because the sites where galls develop already have the predisposition to accumulate more pigment. Here, we call this the host variation hypothesis.
• We test this hypothesis using the system of galls induced by Palaeomystella oligophaga on Macairea radula host plant. Using spectrophotometry, we calculated the Anthocyanin Reflectance Index (ARI) of gall projections, which are responsible for their characteristic colours. We tested the influence of occupant identity (galling insect or any natural enemy), gall volume, parenchyma thickness, height from the ground, ARI of leaf, ARI of gall surface and ARI of the respective stem.
• We corroborated the host variation hypothesis since the anthocyanin content in stems and in galls' projections were positively related. Moreover, anthocyanin in galls' projections was positively related to anthocyanin in the gall surface and negatively related to gall volume and parenchyma thickness. This shows that, besides the host specificities, galls' own traits may also be responsible for pigment accumulation, influencing their colours.
• In this study, using colour as an example, we show that although galls tend to be considered complex expressions of galling insects' stimuli, their traits may be simply influenced by previous and specific attributes of the host organs.

     

Direct and indirect interactions involving ants, insect herbivores, parasitoids, and the host plant Baccharis dracunculifolia (Asteraceae)

Abstract.  1. The relative importance of direct and indirect interactions in controlling organism abundance is still an unresolved question. This study investigated the role of the direct and indirect interactions involving ants, aphids, parasitoids, and the host plant Baccharis dracunculifolia (Asteraceae) on a galling herbivore Baccharopelma dracunculifoliae (Homoptera: Psyllidae).
2. The effects of these interactions on the galling herbivore's performance were evaluated by an exclusion experiment during two consecutive generations of the galling insect.
3. Ants had a direct negative effect on the performance of the galling herbivore by reducing the number of nymphs per gall. In contrast, ants had no indirect effects on gall mortality through the associated parasitoids.
4. Aphids negatively affected gall development, suggesting that galls and aphids might be partitioning photoassimilates and nutrients moving throughout host-plant tissues.
5. In addition, galls that developed during the rainy season were heavier, indicating that variation in the host plant, due to weather changes, can affect the development of B. dracunculifoliae galls. However, variation in the development of B. dracunculifoliae galls due to presence of aphids or the weather changes did not affect parasitoid attack.
6. These results suggest that direct interactions between ants and galls influenced galling insect abundance, whereas numerical indirect effects involving galling insects, ants, aphids, and host plants were less conspicuous.     

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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Interactions among three trophic levels and diversity of parasitoids: a case of top-down processes in Mexican tropical dry forest

The objective of this study was to analyze the relationship between plant hosts, galling insects, and their parasitoids in a tropical dry forest at Chamela-Cuixmala Biosphere Reserve in western Mexico. In 120 transects of 30 by 5 m (60 in deciduous forest and 60 in riparian habitats), 29 galling insects species were found and represented in the following order: Diptera (Cecidomyiidae, which induced the greatest abundance of galls with 22 species; 76%), Homoptera (Psylloidea, 6.9%; Psyllidae, 6.9%; Triozidae, 3.4%), Hymenoptera (Tanaostigmatidae, 3.4%; which were rare), and one unidentified morphospecies (3.4%). In all cases, there was a great specificity between galling insect species and their host plant species; one galling insect species was associated with one specific plant species. In contrast, there was no specificity between parasitoid species and their host galling insect species. Only 11 species of parasitoids were associated with 29 galling insect species represented in the following families: Torymidae (18.2%), Eurytomidae (18.2%), Eulophidae (18.2%), Eupelmidae (9.1%), Pteromalidae (9.1%), family Braconidae (9.1%), Platygastridae (9.1%), and one unidentified (9.1%). Most parasitoid species parasitized several gall species (Torymus sp.: 51.1%, Eurytoma sp.: 49.7%, Torymoides sp.: 46.9%). Therefore, the effects of variation in plant defenses do not extend to the third trophic level, because a few species of parasitoids can determine the community structure and composition of galling insect species in tropical plants, and instead, top-down processes seem to be regulating trophic interactions of galling insect species in tropical gall communities.     

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.     

Evolutionary relationships of Pemphigus and allied genera (Hemiptera: Aphididae: Eriosomatinae) and their primary endosymbiont,Buchnera aphidicola

Aphids harbor primary endosymbionts, Buchnera aphidicola, in specialized cells within their body cavities. Aphids and Buchnera have strict mutualistic relationships in nutrition exchange. This ancient association has received much attention from researchers who are interested in endosymbiotic evolution. Previous studies have found parallel phylogenetic relationships between non‐galling aphids and Buchnera at lower taxonomic levels (genus, species). To understand whether relatively isolated habitats such as galls have effect on the parallel relationships between aphids and Buchnera, the present paper investigated the phylogenetic relationships of gall aphids from Pemphigus and allied genera, which induce pseudo‐galls or galls on Populus spp. (poplar) and Buchnera. The molecular phylogenies inferred from three aphid genes (COI, COII and EF‐1α) and two Buchnera genes (gnd, 16S rRNA gene) indicated significant congruence between aphids and Buchnera at generic as well as interspecific levels. Interestingly, both aphid and Buchnera phylogenies supported three main clades corresponding to the galling locations of aphids, namely leaf, the joint of leaf blade and petiole, and branch of the host plant. The results suggest phylogenetic conservatism of gall characters, which indicates gall characters are more strongly affected by aphid phylogeny, rather than host plants.     

Fossil oak galls preserve ancient multitrophic interactions

Trace fossils of insect feeding have contributed substantially to our understanding of the evolution of insect-plant interactions. The most complex phenotypes of herbivory are galls, whose diagnostic morphologies often allow the identification of the gall inducer. Although fossil insect-induced galls over 300Myr old are known, most are two-dimensional impressions lacking adequate morphological detail either for the precise identification of the causer or for detection of the communities of specialist parasitoids and inquilines inhabiting modern plant galls. Here, we describe the first evidence for such multitrophic associations in Pleistocene fossil galls from the Eemian interglacial (130000-115000 years ago) of The Netherlands. The exceptionally well-preserved fossils can be attributed to extant species of Andricus gallwasps (Hymenoptera: Cynipidae) galling oaks (Quercus), and provide the first fossil evidence of gall attack by herbivorous inquiline gallwasps. Furthermore, phylogenetic placement of one fossil in a lineage showing obligate host plant alternation implies the presence of a second oak species, Quercus cerris, currently unknown from Eemian fossils in northwestern Europe. This contrasts with the southern European native range of Q. cerris in the current interglacial and suggests that gallwasp invasions following human planting of Q. cerris in northern Europe may represent a return to preglacial distribution limits.     

Unexpected High Diversity of Galling Insects in the Amazonian Upper Canopy: The Savanna Out There

A relatively large number of studies reassert the strong relationship between galling insect diversity and extreme hydric and thermal status in some habitats, and an overall pattern of a greater number of galling species in the understory of scleromorphic vegetation. We compared galling insect diversity in the forest canopy and its relationship with tree richness among upland terra firme, várzea, and igapó floodplains in Amazonia, Brazil. The soils of these forest types have highly different hydric and nutritional status. Overall, we examined the upper layer of 1,091 tree crowns. Galling species richness and abundance were higher in terra firme forests compared to várzea and igapó forests. GLM-ANCOVA models revealed that the number of tree species sampled in each forest type was determinant in the gall-forming insect diversity. The ratio between galling insect richness and number of tree species sampled (GIR/TSS ratio) was higher in the terra firme forest and in seasonally flooded igapó, while the várzea presented the lowest GIR/TSS ratio. In this study, we recorded unprecedented values of galling species diversity and abundance per sampling point. The GIR/TSS ratio from várzea was approximately 2.5 times higher than the highest value of this ratio ever reported in the literature. Based on this fact, we ascertained that várzea and igapó floodplain forests (with lower GIA and GIR), together with the speciose terra firme galling community emerge as the gall diversity apex landscape among all biogeographic regions already investigated. Contrary to expectation, our results also support the “harsh environment hypothesis”, and unveil the Amazonian upper canopy as similar to Mediterranean vegetation habitats, hygrothermically stressed environments with leaf temperature at lethal limits and high levels of leaf sclerophylly.     

Host plant effects on the development and survivorship of the galling insect Neopelma baccharidis (Homoptera: Psyllidae)

In this study, the mortality factors acting upon the galling psyllid Neopelma baccharidis Burckhardt (Homoptera) caused by its host plant, Baccharis dracunculifolia De Candole (Asteraceae) were analysed. In March 1999, 982 galls of the same cohort were randomly marked on 109 individuals of B. dracunculifolia in the field. Galls were censused each month during their development, from April to August, and dead galls were collected and analysed for mortality factors. Gall dehiscence rates were calculated for each month. The major mortality source of N. baccharidis was gall dropping (13.2% of the original cohort), which is probably a normal outcome of previous mortality caused by the other factors observed in this study. Unknown factors killed 11.7% of this gall population and were ascribed to plant resistance during gall development. Empty galls represented 7.7% of the observed mortality and may be a consequence of egg retention or egg mortality/abortion related to variations in plant quality. Shoot mortality was high during the dry season and killed 7.5% of the galls, but this impact was minimized after the third month from gall formation due to the ability of nymphs to accelerate development and emerge from galls on dying shoots. However, the size of dehisced galls on dead shoots tended to be smaller, possibly affecting adult performance. Mortality of N. baccharidis attributed to B. dracunculifolia strongly controlled the galling insect population, killing 40.7% of the original cohort of galls. Plant‐mediated mortality was caused by often neglected factors acting predominantly during the first 3 months of development, which are critical to gall survivorship. These results reinforce the importance of bottom‐up forces in plant‐insect systems.     

Fungal endophyte communities differ between chestnut galls and surrounding foliar tissues

Foliar endophytic fungi are present in almost all vascular plants. The composition of endophyte communities varies among plant individuals. Likely, but understudied, sources of this variation are the species composition of the plant community and initial attacks by insect herbivores. We addressed these issues by characterizing fungal endophyte communities on leaves of chestnut (Castanea sativa) grown in pure vs. mixed stands. We used ITS metabarcoding methods to identify endophytic fungi associated with galls caused by the invasive gall wasp, Dryocosmus kuriphilus, and with surrounding chestnut leaf tissues. We found 1378 different OTUs. The richness, diversity and composition of endophyte communities differed between galls and surrounding leaf tissues but were independent of forest stand composition. Fungal endophyte richness was lower in galls than in surrounding leaf tissues. Most differences in the composition of fungal endophyte communities between galls and foliar tissues were due to OTU turnover. These results suggest that insect-induced galls provide a particular habitat condition for endophytic microorganisms, regardless of forest species composition. A better understanding of endophyte biology is important to improve their use as biocontrol agents of galling insects.     

Morphometric analysis of young petiole galls on the narrow-leaf cottonwood, <Emphasis Type="Italic">Populus angustifolia</Emphasis>, by the sugarbeet root aphid, <Emphasis Type="Italic">Pemphigus betae</Emphasis>

An insect-induced gall is a highly specialized structure resulting from atypical development of plant tissue induced by a reaction to the presence and activity of an insect. The insect induces a differentiation of tissues with features and functions of an ectopic organ, providing nutrition and protection to the galling insect from natural enemies and environmental stresses. In this anatomical and cytological study, we characterized how the gall-inducing aphid Pemphigus betae reshapes the leaf morphology of the narrow-leaf cottonwood Populus angustifolia to form a leaf fold gall. Young galls displayed a bend on one side of the midvein toward the center of the leaf and back to create a fold on the abaxial side of the leaf. This fold was formed abaxially by periclinal and anticlinal divisions, effectively eliminating intercellular spaces from the spongy parenchyma. Galls at this stage exhibited both cell hypertrophy and tissue hyperplasia. Cells on the adaxial surface were more numerous and smaller than cells near the abaxial surface were, creating the large fold that surrounds the insect. Mesophyll cells exhibited some features typical of nutritive cells induced by other galling insects, including conspicuous nucleolus, reduced and fragmented vacuole, smaller and degraded chloroplasts, and dense cytoplasm compared to ungalled tissue. Even though aphids feed on the contents of phloem and do not directly consume the gall tissue, they induce changes in the plant vascular system, which lead to nutrient accumulation to support the growing aphid numbers in mature galls.     

Hypersensitivity of Fagus sylvatica L. against leaf galling insects

Hypersensitivity is known as a localized resistance of plants against pathogens. It also can be detected in response to galling insects, i.e., in the area immediately adjacent to the site of oviposition and attempted penetration by the galling larva. This host response includes morphological and histological changes that cause the death of the attacked tissue. It is observed as a rounded dark brown halo around the gall induction site. We provide the first observation on the occurrence and possible relevance of this induced mechanism by which one of the most common tree species in Germany, Fagus sylvatica L., resists attack by two of its most common galling insects, Mikiola fagi and Hartigiola annulipes (Diptera: Cecidomyiidae). Galls induced by these cecidomyiids were extremely common in the studied area in beech forests around Darmstadt, Germany. The availability of resources (leaves on a stem) was a poor predictor of attack by the galling insects as well as for gall abundance (galls successfully formed). Hypersensitive reaction was the most important factor acting against the galling population studied. More than 77% of the attempts of the insects to induce galls on F. sylvatica resulted in failure and consequently the death of the galling larvae. Therefore, few live galls remained to be found and destroyed by natural enemies. This corroborates the view that in galling insect–host plant system interactions plant-driven factors may play a major role in determining herbivore failure and success, and perhaps the resulting community structure.     

The occurrence and effectiveness of hypersensitive reaction against galling herbivores across host taxa

1. Hypersensitive reaction is an important type of induced defence by which the plant elicits a defence response to pathogens and insects. Hypersensitive reaction has been argued to be the most common plant resistance mechanism against insect herbivores that have intimate associations with their host plants. 2. The work reported here attempted to establish how important and widespread hypersensitive reaction might be against gall‐forming species across host taxa. 3. Hypersensitive reaction was the most important mortality factor against gall formation across host plant taxa in seven out of eight cases. 4. The number of insect galls correlated with the size of the leaves but module (leaf) size was a weak factor influencing the incidence of plant hypersensitive reaction to galling. 5. Insect galls and hypersensitive reactions occurred in genetically distant as well as geographically widespread host plant taxa.