Taking a chance on moths: oviposition by Delia flavifrons (Diptera: Anthomyiidae) on the flowers of bladder campion, Silene vulgaris (Caryophyllaceae)

Abstract.

• 1 Delia flavifrons Zetterstedt (Diptera: Anthomyiidae) visits flowers of Silene vulgaris (Moench) Garcke (Caryophyllaceae), where the adults feed, mate, lay their eggs, and the larvae feed on developing seeds. The objective of the study was to examine how an ovipositing female fly assures a food resource for her progeny.
• 2 Ovipositing females preferred young, non-pollinated flowers over older pollinated ones. The flies did not pollinate the flowers and survival of the larvae depended on the flowers being pollinated by moths.
• 3 Flowers containing fly eggs were pollinated more often than expected from chance, probably as a result of both flies and moths visiting particular flowers.
• 4 Eggs were laid singly, and multiple oviposition occurred randomly. Although most eggs hatched, only about half produced larvae that made their way into the fruits. As a result, the probability of competition arising from multiple oviposition may be reduced to such an extent that selection does not favour females that avoid flowers with conspecific eggs.
• 5 Moth larvae of the noctuid genus Hadena also feed on the seed pods of S. vulgaris and will kill any fly larvae they encounter. However, there was no deviation from random oviposition by the flies in relation to eggs laid by the moths, but the competitively weaker fly usually started to lay eggs towards the end of the moth's egg-laying period.
• 6 The relationship between Delia flavifrons and Silene vulgaris superficially parallels that for known pollinator/predator systems, but floral adaptations to hypothetically pollinating flies seems not to have taken place.

     

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.

     

Flower morphology evolution in Jaborosa (Solanaceae): shape and size variation associated with contrasting pollination modes

• Flower morphology is considered an important factor in species diversification because it may influence the efficiency of pollination in different ways (e.g. attraction and mechanical fit with different groups of pollinators). In the present study, we quantified the variation in flower morphology (i.e. shape and size) of the diverse South American genus Jaborosa Juss. (Solanaceae) in relation to contrasting pollination modes: rewarding pollination either by moths or by generalist small insects versus brood-site deceptive pollination by saprophilous flies.
• We examined variations of flower morphology in frontal (pollinator attraction) and sagittal (functional fit with pollinators) views in 12 Jaborosa species using geometric morphometric methods and comparative approaches to infer whether flower shape evolution, not attributable to flower allometry or phylogenetic relationship, is associated with shifts in pollination modes.
• We found remarkable variation in flower morphology among both Jaborosa species and pollination modes, largely in sagittal view. Evolutionary trends in shape of fly-pollinated flowers were mainly attributable to changes in developmental trajectories. Variation in flower architecture facilitated differential pollen placement – on the proboscis of moths, and either on the back or ventral region of saprophilous flies – promoting diversification of the genus.
• Diversification of shape, independent of size, in most of the studied Jaborosa species would indicate adaptation to contrasting pollination modes.

     

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.

     

A plant pathogen causes extensive mortality in an invasive insect herbivore

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Timing of growth determines fitness and performance of a galling insect on willow

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MEASURING SELECTION ON REACTION NORMS: AN EXPLORATION OF THE EUROSTA-SOLIDAGO SYSTEM

The sensitivity of genotypic expression to the environment can be depicted as the reaction norm, which is defined as the array of phenotypes produced by a single genotype over a range of environments. We studied selection on reaction norms of the gall-inducing insect Eurosta solidaginis (Diptera; Tephritidae), which attacks tall goldenrod Solidago altissima (Compositae). Gall size was treated as a component of insect phenotype and attributes of the host plant as environmental influences on gall development. Genetic differences in the response of gall size to plant lag time (the number of days before a plant responds to the gall maker) were examined. Reaction norms for full-sib families of flies were quantified as linear functions; the elevation of the function denoted gall size produced by the family averaged across all plants, and the function's slope denoted family sensitivity to lag time. Expected fitness of each family was regressed over reaction norm elevation and slope to yield selection gradients on these reaction norm parameters. Directional selection on gall size averaged across environments is four times stronger than selection on sensitivity. Yet, genetic variation for sensitivity contributes more than twice as much to gall phenotypic variance as family mean gall size. Our results suggest that selection on environmental sensitivity will be weak for populations restricted to a narrow segment of an environmental gradient, but strong for broadly distributed species.     

Morphological specialization of heterantherous Rhynchanthera grandiflora (Melastomataceae) accommodates pollinator diversity

• The tropical Melastomataceae are characterized by poricidal anthers which constitute a floral filter selecting for buzz‐pollinating bees. Stamens are often dimorphic, sometimes with discernible feeding and pollinating functions. Rhynchanthera grandiflora produces nectarless flowers with four short stamens and one long stamen; all anthers feature a narrow elongation with an upwards facing pore.
• We tested pollen transfer by diverse foraging bees and viability of pollen from both stamen types. The impact of anther morphology on pollen release direction and scattering angle was studied to determine the plant's reproductive strategy.
• Medium‐sized to large bees sonicated flowers in a specific position, and the probability of pollen transfer correlated with bee size even among these legitimate visitors. Small bees acted as pollen thieves or robbers. Anther rostrum and pore morphology serve to direct and focus the pollen jet released by floral sonication towards the pollinator's body. Resulting from the ventral and dorsal positioning of the short and long stamens, respectively, the pollinator's body was widely covered with pollen. This improves the plant's chances of outcrossing, irrespective of which bee body part contacts the stigma. Consequently, R. grandiflora is also able to employ bee species of various sizes as pollen vectors.
• The strategy of spreading pollen all over the pollinator's body is rather cost‐intensive but counterbalanced by ensuring that most of the released pollen is in fact transferred to the bee. Thus, flowers of R. grandiflora illustrate how specialized morphology may serve to improve pollination by a functional group of pollinators.

     

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.

     

Bottom-up effects shift top-down community structure of the parasitoid guild that attacks Asphondylia borrichiae (Diptera: Cecidomyiidae) on Borrichia frutescens (Asteraceae)

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Population biology of swift (Apus apus) ectoparasites in relation to host reproductive success

Abstract.

• 1 We censused ectoparasite populations of adult and nestling swifts over the course of the host's breeding season. Nearly all of the birds were infested with chewing lice and two-thirds of the nests were infested with louse flies. Feather mites were observed but not quantified.
• 2 Lice and louse flies both showed aggregated distributions among hosts. Louse eggs, hatched lice and adult louse flies had negative binomial distributions, whereas the aggregated distribution of louse fly pupae was not adequately described by negative binomial or Poisson models.
• 3 Transmission of lice from parents to offspring was documented. A comparison of the age structure of lice on parents and offspring indicated that most transmission was by nymphal lice.
• 4 Host reproductive success and survival appeared to be independent of the number of lice or louse flies. Neither parasite correlated with the number, body mass, or date of fledging of young birds, nor with the overwinter survival of adults. We caution, however, that experimental manipulations of parasite load are required for a definitive test of the impact of ectoparasites on evolutionary fitness components.

     

Response of Drosophila suzukii (Diptera: Drosophilidae) to extreme heat and dryness

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Competitor‐free space mediates non‐target impact of an introduced biological control agent

Abstract 1. Enemy‐free space has been shown to mediate host shifts in herbivores, but this has not previously been documented in parasitoids. Also, natural enemies shown to maintain host shifts have always been from higher trophic levels, rather than competitors. 2. In Hawaii, an Australian parasitoid (Diachasmimorpha tryoni) of medflies that loses competition contests to a subsequently introduced Asian parasitoid (Fopius arisanus) has shifted its realised host range to attack non‐target gall flies on lantana. 3. The present study demonstrates experimentally that D. tryoni reproduction is: (i) lower on medflies in coffee when F. arisanus is present than when it is absent; (ii) higher in gall flies on lantana than on medflies in coffee, when F. arisanus is present; and (iii) higher in medflies on coffee than in gall flies on lantana, when F. arisanus is absent. This meets Berdegue et al.’s (Ecological Entomology, 21 , 203–217, 1996) three conditions to confirm the importance of enemy‐free space. 4. In the field, F. arisanus is abundant on medflies, whereas D. tryoni is rare on medflies, but is the dominant parasitoid of lantana gall flies. 5. Competitor‐free space is thus shown to be a key mechanism maintaining an apparent host shift by an introduced biocontrol agent onto a non‐target species.     

The life history of a gall‐inducing mite: summer phenology,predation and influence of gall morphology in a sugar maple canopy

• 1 Eriophyoid mites are among the most ubiquitous gall‐inducing arthropods, and are adapted species‐specifically to a broad diversity of plants, although their life histories remain poorly studied outside agricultural systems.
• 2 We examined the seasonal phenology of a leaf‐galling eriophyid mite, the maple spindle gall mite Vasates aceriscrumena (MSGM), in naturally occurring stands of sugar maple Acer saccharum in south‐central Ontario in 2007 and 2008.
• 3 Galls were first induced in spring (mid‐May) and were devoid of mites by late August. In the study region, MSGM appears to have at least two generations, with overwintering, deutogyne females that initiate galls in spring (mid‐May) after leaf flush, giving rise to a generation of protogyne (primary) females and a few morphologically similar males (<1 for every 10 females) and, subsequently, to a new generation of deutogyne females in mid‐July to early August. In July, some galls can be highly crowded, with 50–200 individuals per gall.
• 4 In addition, a tarsonemid mite, Tarsonemus acerbilis, was found in approximately 40% of MSGM galls examined. As much as 95.4% of galls in 2007 and 97.4% in 2008 that contained tarsonemid larvae did not contain MSGM eggs (by contrast, only 2.3% of tarsonemid‐free galls contained no MSGM eggs), suggesting that these juveniles feed, at least opportunistically, on MSGM eggs.
• 5 Gall ostiole morphology appeared to influence both MSGM and Tarsonemus densities within galls, with ‘open’ ostioles (versus ‘closed’) being much more susceptible to invasion by the tarsonemid. The latter is likely to be an important regulator of MSGM populations. We hypothesize that the two ostiole types are the result of selection pressures on the gall inducer, favouring closed gall entrances for increased protection, and possibly also on the host tree, favouring open galls to increase predator access.

     

Alien herbivores and native parasitoids: rapid developments and structure of the parasitoid and inquiline complex in an invading gall wasp Andricus quercuscalicis (Hymenoptera: Cynipidae)

Abstract.

• 1 Rapid and substantial changes have occurred in the parasitoid and inquiline community associated with the agamic galls of Andricus quercuscalicis since it invaded Britain in the late 1950s. The number of parasitoid and inquiline species has risen from one to thirteen over a 15-year period. Although the number of species has been relatively consistent over the last 8 years, the species composition has changed considerably and in a highly characteristic way during this period.
• 2 The parasitoid complex can be divided into two broadly distinct sets of parasitoid species; one set attacks only the gall former whereas the other set concentrates on the inquilines living in the wall of the gall.
• 3 The most dramatic change, however, is in the abundance of inquilines which were reported to be virtually absent in earlier studies on this community in Britain. Over a period of only 5 years, between 1988 and 1993, inquiline attack rose from less than 0.01 to an average of 0.26 inquilines per gall. The intensity of inquiline attack is geographically heterogenous, with high inquiline numbers restricted to south-east England. Because of the relatively high specificity of the parasitoids, high inquiline abundance is positively correlated with parasitoid species richness in knopper galls.
• 4 Parasitism rates, particularly on the gall former, were generally low (<10%). Over the last 5 years, however, seven parasitoid species have been consistently recorded and the mortality caused by these species has increased continuously. The species composition of the community associated with this alien gall wasp in Britain has quickly converged to the community known from its native range in continental Europe. Parasitoid species known to attack the galls of A.quercuscalisis on the continent have been recorded from it in Britain for the first time mainly in areas where inquilines have recently become abundant.
• 5 Since rates of parasitism of the gall former are still low, parasitoids are unlikely to play a major role in the population dynamics of this invading gall wasp at present, but the rapidly increasing inquiline and parasitoid attack could be a source of increased mortality for native cynipid species which are the alternative hosts of those parasitoid species.

     

Coastal insect herbivore populations are strongly influenced by environmental variation

Abstract.

• 1 Pissonotus quadripustulatus is a brachypterous planthopper that feeds on the clonal salt marsh plant, sea oxeye daisy, Borrichia frutescens. Asphondylia borrichiae is a macropterous gall fly that creates galls on the apical meristems. Most Borrichia occurs in isolated patches consisting of more than one genotype.
• 2 Densities of Pissonotus and Asphondylia do not significantly differ within patches but they do differ between patches. I tried to find out why some patches of Borrichia support consistently higher densities of Pissonotus and Asphondylia than others.
• 3 Reciprocal transplants of Borrichia between patches showed that for both Pissonotus and Asphondylia recipient site (local environment) had a strong effect on population densities. However, the best sites for Asphondylia were generally the worst sites for Pissonotus and vice versa.
• 4 Donor site (local plant genotypes) was not significant for Pissonotus but was marginally significant for Asphondylia. However, for Asphondylia recipient site had a greater effect. Local plant genotype effects may be greater for gall-forming insects, which tend to be specialists, than for more generalist sapsucking insects.

     

Preference induction and the benefits of floral resources for a facultative florivore

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Diversity in Eucalyptus susceptibility to the gall‐forming wasp Leptocybe invasa

• 1 Extensive variation to damage by the invasive gall‐forming wasp Leptocybe invasa Fisher & LaSalle (Hymenoptera: Eulophidae) is known to exist amongst Eucalyptus genotypes.
• 2 In the present study, 30 of the 50 tested genotypes were susceptible to gall formation and development of the wasp. Gall development on the petiole and leaves of plants was compared to calculate the percentage of infestation per plant and per genotype.
• 3 A positive correlation between galls on petioles and leaves indicated an absence of specificity at this level, and also that either leaves or petioles could be used to obtain an accurate estimate of the level of infestation.
• 4 Genotypes of E. nitens×E. grandis and E. grandis×E. camaldulensis were most susceptible, with a maximum damage index value for leaves and petioles of 0.52 and 0.39, respectively. Eucalyptus dunii, E. nitens, E. smithii, E. urophylla and E. saligna×E. urophylla showed little or no infestation.
• 5 The results obtained in the present study suggest that the selection and planting of resistant/less susceptible genotypes will be an important aid in managing damage from L. invasa invasion.

     

Use of forested field edges by a blueberry insect pest,Rhagoletis mendax (Diptera: Tephritidae)

1. The blueberry maggot fly, Rhagoletis mendax Curran, is a pest of wild and highbush blueberries. In wild blueberries, most flies colonize fields from forest edges annually. Flies were associated with forest canopies adjacent to colonized fields.
2. Flies were captured up to 6 m high (limited by trap placement height) and may occur at greater tree heights. Fly abundance relative to tree species was highest in red oak.
3. Releases of marked flies from tree canopies showed that release height had no effect on distance into the field that flies dispersed. This is important because the main fly control tactic is the application of a 25- to 30-m-wide swath of insecticide along field perimeters.
4. In the laboratory, flies preferred leaves and leaf extracts of red oak compared to other leaf tree and shrub species, including blueberry. Electroantennographs demonstrated that the female antennae consistently responded to compounds extracted from red oak leaves. Mass spectra and gas chromatograph retention times of five antenna-active compounds matched those of (i) trans-β-ocimene, (ii) linalool, (iii) 4,8-dimethyl-1,3,7-nonatriene, (iv) indole, and (v) trans-nerolidol.
5. We speculate flies recruit to trees before movement into blueberry fields primarily for food, as supported by their arrestment response to leaf extracts.

     

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.