A gall-inducing arthropod drives declines in canopy tree photosynthesis

Mature forest canopies sustain an enormous diversity of herbivorous arthropods; however, with the exception of species that exhibit large-scale outbreaks, canopy arthropods are thought to have relatively little influence on overall forest productivity. Diminutive gall-inducing mites (Acari; Eriophyoidae) are ubiquitous in forest canopies and are almost always highly host specific, but despite their pervasive occurrence, the impacts of these obligate parasites on canopy physiology have not been examined. We have documented large declines in photosynthetic capacity (approx. 60%) and stomatal conductance (approx. 50%) in canopy leaves of mature sugar maple (Acer saccharum) trees frequently galled by the maple spindle gall mite Vasates aceriscrumena. Remarkably, such large impacts occurred at very low levels of galling, with the presence of only a few galls (occupying approx. 1% of leaf area) compromising gas-exchange across the entire leaf. In contrast to these extreme impacts on the leaves of adult trees, galls had no detectible effect on the gas-exchange of maple saplings, implying large ontogenetic differences in host tolerance to mite galling. We also found a significant negative correlation between canopy tree radial increment growth and levels of mite galling. Increased galling levels and higher physiological susceptibility in older canopy trees thus suggest that gall-inducing mites may be major drivers of “age-dependent” reductions in the physiological performance and growth of older trees.     

Astegopteryx spinocephala (Hemiptera: Aphididae), a new aphid species producing sterile soldiers that guard eggs laid in their gall

Astegopteryx spinocephala sp. nov., a soldier‐producing aphid species forming banana‐bunch shaped galls on Styrax benzoides in northern Thailand, is described. We found that galls of the species are formed in approximately June and last for almost 1 year, and that the aphid completes its life cycle without migrating to secondary hosts. Many alate sexuparae appeared in March/April, when many sexuals (arostrate males and rostrate females) and eggs were found in live subgalls guarded by soldiers that plugged the ostiole with their sclerotized, spiny heads. Two healthy galls sampled in April contained 2799 and 2659 eggs, respectively. Many live galls were still found in May, and some at the beginning of June. These galls contained both active soldiers and eggs, some of which had already hatched. This indicates that soldiers of A. spinocephala guard eggs until they hatch in at least some galls.     

The use of volatile cues in recognition of kin eggs by predatory mites

1. Several animal species are known to distinguish between their own eggs and eggs of unrelated conspecifics. However, the cues involved in this discrimination are often unknown. These cues were studied using the predatory mite Gynaeseius liturivorus Ehara.
2. Adult females of these predatory mites oviposit in clusters and avoid oviposition close to eggs laid by other females, resulting in reduced cannibalism between offspring. Because predatory mites are blind, it was tested whether volatiles of eggs were used as a cue for egg recognition.
3. Adult female predatory mites were offered volatile cues of their own eggs and of unrelated conspecific eggs, and females were prevented from contacting the eggs. Predatory mites oviposited closer to their own eggs than to unrelated eggs. This preference was observed even when one own and one unrelated egg were offered as a volatile source.
4. These results suggest that adult female predatory mites can determine kinship using volatiles released from the eggs.

     

Galling and reversion disease incidence in a range of blackcurrant genotypes, differing in resistance to the blackcurrant gall mite (Cecidophyopsis ribis) and blackcurrant reversion disease

An assessment was made of the response of 10 blackcurrant genotypes, differing in resistance to the agent of blackcurrant reversion disease (BRD) and to its gall mite vector, in field trials in Scotland and Finland over 5 yr. At each location, the 10 genotypes were planted in plots containing infector plants with high inoculum levels of the two organisms. In Scotland, the infector plants contained large numbers of gall mites (Cecidophyopsis ribis) and were infected with the European (E) form of BRD; in Finland, infector plants contained a different species of gall mite (C. spicata) and the severe Russian form (R) of BRD. As expected, at both sites, almost all plants of cvs Ben Alder, Ben Lomond, Ben Tirran, Ojebyn and an SCRI selection F4/1/66, which are susceptible to gall mite and BRD, became infested with each of these organisms. However, in Scotland but not in Finland, 58% of cv. Ojebyn plants were affected by BRD. The cv. Foxendown, which contains gene Ce that confers apparent immunity to C. ribis, was free from galls and failed to develop distinctive BRD symptoms at both sites. The cvs Rus and Neosypajuscajaija, which contain gene P which is reported to confer resistance to C. ribis, were infested more slowly by mites than the mite-susceptible genotypes and showed a smaller number of galls per plant. Also, they were infected with BRD more slowly than some mite-susceptible genotypes although by the end of the experiment, most plants were affected by BRD. All plants of cvs Golubka and Ben Gairn, which are resistant to the agent of BRD, remained free from distinct BRD symptoms at both sites despite the fact that plants contained galls. These data indicate the superiority of gene Ce over gene P for resistance to gall mites with the added benefit that the virtual immunity to gall mites provided by gene Ce provides a high level of protection against infection with BRD. The relative merits of these different forms of resistance to gall mites and BRD in blackcurrant are discussed in relation to present control methods for these two organisms and in the light of recent findings of different species of eriophyid mites on Ribes species.     

Gall mite Fragariocoptes setiger (Eriophyoidea) changes leaf developmental program and regulates gene expression in the leaf tissues of Fragaria viridis (Rosaceae)

The interaction of plants with certain types of parasites leads to the formation of galls, organised structures that create the habitat of the parasite, caused by an abnormal proliferation of host plant's cells under the influence of growth regulators, secreted by the parasite, or by the plant itself under the influence of the parasite. Arthropods, mites in particular, are the largest group of gall‐inducing phytoparasites, but the mechanisms of their interaction with plants remain virtually unexplored. The interaction of the gall‐inducing eriophyoid mite Fragariocoptes setiger with Fragaria viridis plants was used as a model gall–mite system where data were obtained on the changes in the histological structure of F. viridis leaf blades under the influence of the mites as well as F. viridis gene expression during gall formation. For histological purposes, gall formation was split into four stages with each corresponding to the age of the gall as well as to specific changes that occur during that period. A dramatic change of adaxial–abaxial polarity of the lamina throughout the four stages was observed. Moreover, qRT‐PCR analysis of F. viridis gene expression in the developing gall revealed changes in the expression levels of certain meristem‐specific genes, as well as the genes that determine adaxial–abaxial polarity and signalling of phytohormones.     

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.     

Heat tolerance of strawberry tarsonemid mite Phytonemus pallidus

The heat tolerance of the strawberry tarsonemid mite Phytonemus pallidus ssp. fragariae was studied in northern Sweden by treating adult mites in hot water at temperatures 44°C, 46°C and 48°C. During the summer, the shortest exposure times required to kill all adult mites at these temperatures were 6, 2.5 and 1.25 min, respectively. It is concluded that hot‐water treatment of strawberry runners for 6.5 min at 46°C would suffice to kill all tarsonemid mites on infested runners. This exposure time includes the time needed to raise the temperature of the plant material to that of the water bath. In the autumn, from the middle of September and later, the mites became successively much more heat tolerant as they prepared for hibernation. It is expected that longer exposure periods are needed to kill mites on frigoplants.     

Asian chestnut gall wasp in Tuscany: gall characteristics,egg distribution and chestnut cultivar susceptibility

• 1 Preliminary investigations were carried out on Dryocosmus kuriphilus Yasumatsu on Castanea sativa Miller in Tuscany to assess variations in gall characteristics in coppice and high forest at two crown heights (height < 2 or 2–6 m), influence of bud size and bud position on oviposition rates and susceptibility of three cultivars.
• 2 Gall size may depend on various factors, including wasp population density. In the present study area, small galls (with one or two cells) were the most numerous in 2008, whereas larger galls (with more than three cells) prevailed in 2009.
• 3 Dryocosmus kuriphilus oviposition occurrence was influenced by both bud size and bud position. Buds with eggs tended to be larger in size compared with bud without eggs, suggesting that D. kuriphilus females prefer to lay eggs in larger buds (approximately 6 mm³) compared with smaller buds (approximately 3 mm³). The mean number of eggs per bud tended to decrease from the apical bud toward the basal bud.
• 4 Three C. sativa cultivars, Carpinese, Fusca and Cesurone, were examined. Fusca grafts had significantly more galls compared with Carpinese and Cesurone, whereas Cesurone grafts had more larvae per bud compared with Carpinese and Fusca. Overall, the Carpinese cultivar may be less susceptible to D. kuriphilus galling compared with the Fusca and Cesurone cultivars.

     

Plant-mediated competition facilitates a phoretic association between a gall mite and a psyllid vector

Phoretic associations between mites and insects commonly occur in patchy and ephemeral habitats. As plants provide stable habitats for herbivores, herbivorous mites are rarely dependent on other animals for phoretic dispersal. However, a phoretic gall mite, Aceria pallida, which is found on plants, seasonally attaches to a herbivorous insect, Bactericera gobica, for overwintering survival. After detachment, the gall mite shares a habitat with its vector and is likely to compete with this vector for plant resources. However, excessive competition works against the sustainability of the seasonal phoretic association. How the gall mite, as an obligate phoretic mite, balances this relationship with its vector during the growing season to achieve phoresy is unknown. Here, the plant-mediated interspecific interaction between the gall mite and the psyllid after detachment was studied in the laboratory and field. The laboratory results showed that infestation by the gall mite had detrimental effects on the survival and development of psyllid nymphs. Meanwhile, the mite population and the gall size were also adversely affected. The results from the field showed that the mean densities of the mite galls and psyllids were lower in the mixed-species infestation treatment than in the single-species infestation treatment across the investigation period. However, the interspecific interaction between the gall mite and the psyllid decreased rather than accelerated leaf abscission caused by the psyllid, which promoted the persistence of the psyllid population and then indirectly contributed to phoretic association. Our results suggest that the plant-mediated competition between the phoretic gall mite and its vector after detachment facilitates the maintenance of the phoretic association.     

Characteristics of development and reproduction in Typhlodromus pyri on Tetranychus urticae and Cecidophyopsis ribis. II. Progeny of overwintered females

The results of this study on the bionomic parameters in progeny of overwintered females of Typhlodromus pyri Scheuten under constant laboratory conditions (18°C, R.H. 75%. 18L:6D) are described. The predatory mites were fed overwintering forms of two prey species, the two-spotted spider mite. Tetranychus urticae Koch and the black currant gall mite. Cecidophyopsis ribis (Westwood). The results corroborate that T. pyri is able to complete its life-cycle and reproduce when it feeds exclusively on the black currant gall mite. Juvenile development of females feeding on T. urticae and on C. ribis was 19.4 and 20.9 days respectively, while in males it was 18.4 and 17.5 days. Mortality of juveniles feeding on T. urticae was higher (33.0%) compared with that of juveniles fed on C. ribis (10.7%). The highest mortality was noted in protonymphs fed on T. urticae (21.9%). The difference between the mean total fecundity of females fed on T. urticae (16.4 eggs) and on C. ribis (17.8 eggs) was not statistically significant. Mean duration of preoviposition, oviposition and postoviposition periods were 19.1, 25.0 and 14.5 days in females fed on T. urticae and 19.7. 50.2 and 5.0 in those fed on C. ribis.     

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

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Sexually transmitted disease in a promiscuous insect, Adalia bipunctata

Abstract.

• 1 Sexually transmitted diseases (STDs) have rarely been reported in insects and other invertebrates. The majority of those reported involve organisms where sexual transmission is augmented by either vertical (i.e. inherited) transmission, or horizontal transmission, independent of host sexual activity.
• 2 We here demonstrate the existence of an STD in the coccinellid beetle Adalia bipunctata. This species bears a parasitic mite of the genus Coccipolipus. We show that, like many other podapolipid mites, this mite is transmitted between host individuals at a high rate during copulation. It also appears to be transmitted at a low rate between non-copulating individuals.
• 3 We show that infected female A.bipunctata produced eggs at a reduced rate, and that the eggs produced by infected females have highly decreased viability. However, no effect of infection upon host longevity was observed.
• 4 The results are discussed in relation to the incidences of sexually transmitted disease in invertebrates in general, the causes of disease symptoms, and the importance of this disease in the evolution of A.bipunctata.

     

Life history strategy and taxonomic position of gall midges (Diptera: Cecidomyiidae) inducing leaf galls on Styrax japonicus (Styracaceae)

Four gall midge species (Diptera: Cecidomyiidae) that induce leaf galls on Styrax japonicus (Styracaceae) were identified to generic level based on larval morphology. Three of these gall midges, which induce whitish hemiglobular galls, flattened subglobular galls, and purple globular galls, respectively, were identified as three genetically distinct species of Contarinia, and the remaining species, which induces globular galls with dense whitish hairs, was identified as a species of Dasineura. Field surveys in Fukuoka, Japan, revealed that adults of these gall midges emerged and oviposited in late March to mid‐April at Mount Tachibana (approximately 200 m a.s.l.) and in late April to early May at Mount Sefuri (about 1050 m a.s.l.), coinciding with the leaf‐opening season of S. japonicus. Larvae of these gall midges mostly developed into third instars by June and then left their galls and dropped to the ground. These species therefore have a life history strategy that differs from that of another S. japonicus‐associated gall midge, Oxycephalomyia styraci, which overwinters as the first instar in ovate swellings, matures rapidly in spring, and emerges directly from the galls.     

Role of two phoretic mites in transmission of bluestain fungus, Ceratocystis minor

ABSTRACT.

• 1 Tarsonemus mites phoretic on southern pine beetles were shown to be hypervectors of bluestain fungus, Ceratocystis minor (Hedgcock 1906).
• 2 Beetles with phoretic mites had significantly more C.minor than did mite-free beetles.
• 3 Over 90% of tarsonemid mites collected from beetle-infested inner bark and 59% of mites phoretic on emerging beetles carried C.minor spores.
• 4 This association is an interesting example of a three-way mutualism among insects, mites and fungi.

     

Host plant response and phenotypic plasticity of a galling weevil (Collabismus clitellae: Curculionidae)

The response of a host plant to gall‐inducing insects varies both among and within plants, so that different levels of resources are available to the insects. The weevil Collabismus clitellae Boheman induces galls on the shoots of Solanum lycocarpum St Hil. in south‐east Brazil. Galls are found on a range of parts within an individual plant and are more abundant on smaller plants. In the present study, the host plant response as a possible influence on the performance of C. clitellae both between and within plants was tested. Gall abortion increased with plant height. Within plants, gall size was positively related to shoot diameter and number of chambers within the gall. The increase in gall larval density (number of individuals per gall volume unit) resulted in smaller adults and reduced developmental rates, probably because of resource limitation within the gall. The number of eggs laid by females increased with shoot diameter. Females laid more eggs on thicker shoots, where there are fewer chances to form galls with high larval density. However, this relationship was weak and a large variation was found for adult sizes. The availability of high quality sites is limited to smaller plants and thicker shoots located on the basal region of the plant. The phenotypic plasticity of this insect species in adult size and development time allows individuals growing on low quality sites to reach maturity, thus enhancing exploitation of the host plant.     

Colonization of vineyards by Kampimodromus aberrans (Oudemans) (Acari: Phytoseiidae): dispersal from surrounding plants as indicated by random amplified polymorphism DNA typing

Abstract

• 1 Kampimodromus aberrans (Oudemans) is the most important predatory mite found in vineyards of southern France. This mite also occurs in surrounding uncultivated areas from where it disperses to colonize adjacent vineyards
• 2 To determine accurately origins of immigrant mites and to study their establishment in vineyards, a study using RAPD markers (Random Amplified Polymorphism DNA) was performed. Females of K. aberrans were sampled on leaves collected in an experimental vineyard, and from several adjacent areas including neighbouring vine plots and natural plants, both of which harboured high densities of the mite. Samples were taken in May and July before and after major dispersal of K. aberrans into the experimental plot occurred.
• 3 For both dates, genetic distances within population were lower than between populations and three groupings of mites were observed. Strong relationships were observed between (1) females from different parts of the same experimental vineyard (variety Cabernet‐Sauvignon), (2) females from several plants in the woody margin neighbouring this experimental plot, and (3) females from two neighbouring vineyards (variety Carignan). Populations seemed to be structured and no correlation between genetic and geographical distances was observed. Hence, definitive conclusions about origins of migrants were not possible.
• 4 Once in a vineyard, mites are probably exposed to selection pressures (i.e. pesticide applications or vine variety characteristics) that largely determine differentiation of populations. Thus, despite many immigrants moving into vineyards, our study indicated that there was limited survival or reproduction of immigrants. Further studies of within vineyard selection factors and impacts on immigrant mites are needed to determine the influence of natural colonization on grape pest management.

     

Effects of large‐scale host plant addition and removal on parasitoid‐mediated associational resistance in the gall midge Asphondylia borrichiae

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Quantitative differences detected in the histology of galls induced by the same aphid species in different varieties of the same host

• Plant galls are abnormal growths caused by an inducer that determines their morphology and anatomy. We qualitatively and quantitatively compared the histological anatomy of five aphid species (Paracletus cimiciformis, Forda marginata, Forda formicaria, Baizongia pistaciae and Geoica wertheimae) that induce galls in Pistacia terebinthus shrubs growing in Israel. We also quantitatively compared these galls to those that the aphids create on the same host in Spain.
• Histological study was conducted following methods described previously by the authors.
• Quantitative differences among the galls were found in five of 12 common anatomical traits: gall thickness, stomatal number in the epidermis‐air, size of vascular bundles, distance of phloem ducts from the lumen and number of intraphloematic schizogenous ducts. Other structures were particular to one or some species: number of cracks in the epidermis–lumen, a sclereid layer, trichomes and microcrystal inclusions. Fisher's tests of combined probabilities showed that the galls induced in Israel were statistically different from those in Spain. In particular, the number of intraphloematic schizogenous ducts was higher in the galls induced in P. terebinthus in Israel. Such differences were also found in other traits related to defence of the gall inhabitant.
• In conclusion, while the gall shape and size are determined mainly by the cecidogenic insect, it seems that the host plant also plays an important role in determining the number/size of quantitative traits, in this case mainly protective structures.

     

Characteristics of development and reproduction in Typhlodromus pyri on Tetranychus urticae and Cecidophyopsis ribis. I. Overwintered females

The results of a study of bionomic parameters in overwintered females of Typhlodromus pyri Scheuten under constant laboratory conditions (18±1°C, RH 75%, 18L:6D) are described. The predatory mites were fed two prey species, the two-spotted spider mite, Tetranychus urticae Koch and the eriophyid gall mite, Cecidophyopsis ribis (Westwood). The results indicate that T. pyri females survive and reproduce on C. ribis as well as on T. urticae. However, the average fecundity of females feeding on C. ribis was lower (13.79 eggs) compared with that of females fed T. urticae (29 eggs). The length of preoviposition period varied greatly (8 to 96 days), the number of ovipositing females increasing at a statistically significant rate on T. urticae. Differences in the mean duration of postoviposition period (35.27 and 52.50 days) and longevity in the laboratory (97.82 and 116.04 days) were not statistically significant. Accumulation of metabolites in the alimentary canal was noted in some females of T. pyri.     

A plant pathogen causes extensive mortality in an invasive insect herbivore

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