Effects of urbanisation on the parasitoid community of a leafminer

Urbanisation may have detrimental effects on communities of parasitoids, affecting their species richness, abundance, and species dominance. Here we investigated the influence of the degree of urbanisation on parasitoid communities of Liriomyza commelinae (Frost) (Diptera: Agromyzidae), a leafminer of Commelina erecta L. (Commelinaceae), in the city of Córdoba, Argentina. To study changes in species richness, the specific composition of parasitic complexes and their degree of impact on the leafminer, 18 sampling sites from the centre to the outskirts of the city were selected and different variables indicative of urbanisation were quantified in each site. During January and February of 2005 and 2006, all mined leaves found in each plant patch were collected and the following variables were estimated: proportion of mined patches, abundance of the leafminer, total parasitoid species richness, total parasitism rates and parasitism due to the most abundant parasitoid species. The percentage of mined patches and leafminer abundance increased with urbanisation degree. Estimates of parasitoid species richness were not influenced by urbanisation degree but increased with species richness of mined plants. Changes in the specific composition of species along the urbanisation gradient were observed. Although parasitism by one of the species studied was higher in more urbanised sites of the city, the total parasitism rate of L. commelinae was not affected by urbanisation degree, species richness of mined plants or leafminer abundance. It appears that urbanisation benefits the herbivore species here studied but not through altering parasitoid activity. Changes in parasitoid community composition reflex dissimilar tolerance to environmental conditions displayed by different parasitoid species.     

Quantitative food webs of lepidopteran leafminers and their parasitoids in a Japanese deciduous forest

Quantitative food webs were constructed to explore the community structure of leaf-mining moths in the family Gracillariidae and their parasitoid wasps in a deciduous forest in Hokkaido, Japan. A whole food web was constructed from data collected from June to October 2001. In the web, 16 leafminer species on seven tree species were attacked by 58 species of hymenopteran parasitoid; 376 links between leafminers and parasitoids were observed. Leafminers were specialist herbivores, but most parasitoids were generalists. Five webs were constructed for the seasonal prevalence of leafminers over the one-year period to reveal the temporal dynamics in community structure. Among the seasonal webs, the first web in June was distinctive because two tree species, Japanese umbrella tree Magnolia obovata and Japanese magnolia M. kobus, supported the community. Second to fourth webs from July to September were dominated by the leafminer species on Japanese oak Quercus crispula, and the fifth web was marked by that on Carpinus cordata. The extent of potential apparent competition among leafminers was evaluated using quantitative parasitoid overlap diagrams. These diagrams suggested that abundant host species are likely to have large indirect effects on less abundant species. Moreover, the potential for apparent competition between leafminer species inhabiting different host tree species can occur, although leafminers sharing the same tree species are prone to interact via shared parasitoids. In this system, particular leafminer species, acting as potential sources of apparent competition, can affect other species as sinks, and control whole-community dynamics. Directed apparent competition may potentially occur around oak trees.     

Historical incidence of Perthida leafminer species (Lepidoptera) in southwest Western Australia based on herbarium specimens

Examination of more than 500 herbarium specimens of Eucalyptus marginata and E. rudis for the presence of two species of leafminers, Perthida spp., indicated that these native insect species were very rare in southwest Western Australia until first recorded in metropolitan Perth in 1878 and 1897, respectively, when west coast populations of both leafminer species evidently increased dramatically. The first record from the south coast dates from 1901 near Albany. Evidence of leafminer populations on E. marginata remote from coastal settlements was not recorded until 1917, near Cranbrook. The first record in jarrah forest was in 1967, some 10 years after outbreaks of the pest insect are known to have occurred in the forest. The Perthida leafminer species on E. rudis was first recorded from inland areas in 1904 and appears to have increased more rapidly in abundance than the species found on E. marginata. These early geographical and species differences in incidence are discussed in terms of changing disturbance patterns, which have not been simultaneous across southwest Western Australia.     

Parasitoids as a mortality factor for the lime leafminer (<Emphasis Type="Italic">Phyllonorycter issikii</Emphasis>, Lepidoptera,Gracillariidae)

The species structure and dynamics of the parasitoid complex of the lime leafminer Phyllonorycter issikii Kumata (Lepidoptera, Gracillariidae) were studied in three test plots in Izhevsk (Udmurtia) during 2001–2005. The leafminer mortality rate associated with activity of this complex was assessed.     

Host range and genetic strains of leafminer flies (Diptera: Agromyzidae) in eastern Brazil reveal a new divergent clade of Liriomyza sativae

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Parasitism of different larval stages of Cameraria ohridella

About 20 species of parasitic Hymenoptera havebeen reported from the horse chestnutleafminer, Cameraria ohridella Deschka & Dimic (Lepidoptera: Gracillariidae). Generally,parasitism is low compared to other closelyrelated leafminers and the parasitism levelvaries considerably. Among other reasons,parasitism rates depend on the developmentalstage of the moth and therefore vary with thesampling date. In the current study,investigations on the parasitism of the moth'sfirst generation were carried out in order todetermine which preimaginal stages areparasitized by the most abundant parasiticwasps. Minotetrastichus frontalis (Nees),Pnigalio agraules (Walker) and Chrysocharis nephereus (Walker) (allHymenoptera: Eulophidae) developed as larval orpupal parasitoids and preferred later larvalinstars of the moth. Egg parasitism did notoccur. Overall, the last two of the six larvalinstars (the spinning instars) of the leafminersuffered the heaviest attack. In concordancewith the fairly low parasitism rates, theeffect of the four most abundant chalcidoids onthe leafminer population was negligible, havingno significant influence on the mortality ofC. ohridella. Although the speciescomposition of the parasitoid complex of C. ohridella shows similarities with thesituation found in other closely relatedleafmining moths, it is unlikely that thenaturally occurring chalcidoids will be able toprevent the horse chestnut leafminer fromdeveloping epidemic population densities in thenear future.     

Native parasitoids and their potential to control the invasive leafminer, Cameraria ohridella DESCH. & DIM. (Lep.: Gracillariidae)

In spite of the fact that since the end of the eighties, the horse chestnut leafminer, Cameraria ohridella, has established itself throughout Europe, native predators such as ants and birds are not attuned to this neozoic species. In contrast, several parasitic wasp species already started to exploit the invasive horse chestnut leafminer, but until now parasitation rates are quite low, mainly because of asynchrony in the lifecycles of parasitoids and host. Only the removal of leaf litter, in which pupae hibernate, is at the moment a strategy to reduce the infestation level in the next year. Unfortunately, not only hibernating horse chestnut leafminers but also parasitoids are removed, and important resources for biocontrol are unused. In the current study, we investigated the potential efficiency of the horse chestnut leafminer parasitoid complex extracted from leaf litter in defined environments. Parasitoids were released at different densities to investigate density dependence in parasitation rates.Although seven different species were released in our experiments, only Pnigalio agraules turned out to be responsible for biocontrol of C. ohridella. We recorded parasitation rates of up to 35%. Overall, parasitation rates were independent of the leafminer density but increased fourfold if ten times more parasitoid individuals were released. Unfortunately, none of the parasitoid species could be established in the experimental units in the long run. Results are compared to other parasitoid-leafminer systems, and promotion of horse chestnut leafminer parasitoids to support natural selection and biological control of the horse chestnut leafminer is discussed.     

Impact of natural enemies on coffee leafminer Leucoptera coffeella (Lepidoptera: Lyonetiidae) population dynamics in Chiapas, Mexico

Coffee leafminer, Leucoptera coffeella, is a pest in many New World coffee growing areas. Previous studies suggested that its population dynamics were strongly affected by natural enemies, particularly of larvae, and physical environmental conditions. Our study documented through field surveys and life table analyses (i) the natural enemy complex associated with coffee leafminer and (ii) the impacts of natural enemies on the population dynamics of coffee leafminer, on coffee (Coffea arabica) at two elevations and two rainfall levels in the Soconusco region of Chiapas, Mexico. Twenty-two larval parasitoid species (including 14 morphospecies) were collected. Egg and pupal parasitoids were not recovered. Life table analyses showed that parasitism contributed 10% of real mortality, and parasitism rates were 8–10-fold higher at the low (<550 m) versus high (>950 m) elevation; parasitism rates were similar under low (<100 mm) and high (>400 mm) rainfall. Seventeen predator species (including five morphospecies) were collected, of which most were ants (Formicidae, 14 species) that contributed >58% of real mortality. Life table analyses showed that predation rates were higher at high versus low elevation and under high versus low rainfall. Independently of elevation and rainfall, egg predation (likely by ants) was the most important source of indispensable mortality (range = 0.13–0.30), except at low elevation and high rainfall where pupal predation (=0.14) was similarly important. Also, predation was the main source of coffee leafminer larval and pupal mortality during a 13-month period in a low elevation coffee farm and was highest during the rainy season (>400 mm rainfall/month), when coffee leafminer prevalence was highest. Overall, predation of eggs and pupae (the latter particularly at low elevation), mostly by ants, were the most important sources of coffee leafminer mortality. Because ants were the main source of coffee leafminer egg and pupal mortality, their importance and potential role in coffee pest management strategies were discussed.     

A tritrophic analysis of host preference and performance in a polyphagous leafminer

The optimal oviposition theory predicts that oviposition preferences of phytophagous insects should correlate with host suitability for their offspring. As plant host suitability depends not only on its quality as food, but also on its provision of enemy‐free space, we examined the relationship between adult host preference and offspring performance for the leafminer Liriomyza huidobrensis (Blanchard) (Diptera: Agromyzidae) on various host plants, considering also the interaction with natural enemies. Preference and offspring performance were assessed through observational field data and laboratory experiments in central Argentina. Field data suggested a positive host preference – performance linkage, as the leafminer attained larger body size on the crops where it was more abundant. Laboratory trials supported these results: Vicia faba L. (Fabaceae) was the preferred host in the laboratory as well as in the field, performance of L. huidobrensis being also best on this host, with highest survival rates and shortest development time. The actively feeding larval stage showed the largest plant‐related effects. Higher overall parasitism rates were found on plants from which smaller leafminers were reared, reinforcing the preference–performance linkage. On the other hand, the main parasitoid Phaedrotoma scabriventris Nixon (Hymenoptera: Braconidae) reached larger body size, and caused higher mortality rates on crops where the leafminer was larger. Changes in abundance of particular parasitoid species could thus modify overall parasitism trends.     

Variations in the presence of chloride cells in the gills of lampreys (Petromyzontiformes) and their evolutionary implications

Although confined to fresh water, non‐parasitic species of lampreys and the landlocked parasitic sea lamprey, all of which were derived relatively recently from anadromous ancestors, still develop chloride cells, whose function in their ancestors was for osmoregulation in marine waters during the adult parasitic phase. In contrast, such cells are not developed by the non‐parasitic least brook lamprey Lampetra aepyptera, which has been separated from its ancestor for >2 million years, nor by the freshwater parasitic species of the genus Ichthyomyzon. The length of time that a non‐parasitic species or landlocked parasitic form or species has spent in fresh water is thus considered the overriding factor determining whether chloride cells are developed by those lampreys.     

Host range of the European leaf sheath mining midge,Lasioptera donacis Coutin (Diptera: Cecidomyiidae), a biological control of giant reed,Arundo donax L.

The fundamental host range of the arundo leafminer, Lasioptera donacis a candidate agent for the invasive weed, Arundo donax was evaluated. L. donacis collects and inserts spores of a saprophytic fungus, Arthrinium arundinis, during oviposition. Larvae feed and develop in the decomposing leaf sheath channel tissue. Thirty-six closely related and economic grass species along with several key habitat associates were evaluated in no-choice tests. L. donacis and its associated saprophyte completed development only on A. donax, in concurrence with published reports from its native range in Mediterranean Europe. The arundo leafminer feeding leads to premature defoliation, constituting a different mode of attack on the host plant as compared to two previously released insects, the arundo wasp and arundo scale, which feed on shoot tips and rhizomes, respectively. Defoliation of A. donax is expected to increase light penetration into stands of A. donax which increases visibility for law enforcement, reduces the survival of cattle fever ticks, and enhance recovery of the native riparian vegetation along the Rio Grande and other habitats where this weed is invasive.     

Parasitism of Cameraria ohridella (Lepidoptera, Gracillariidae) in natural and artificial horse-chestnut stands in the Balkans

1 The horse‐chestnut leafminer, Cameraria ohridella Deschka & Dimic, was discovered during a mass outbreak on planted horse‐chestnut trees, Aesculus hippocastanum, near Lake Ohrid in Macedonia. However, several peculiarities in the leafminer's biology and ecology, such as its inefficient control through natural enemies, indicate that it may be of exotic origin. 2 In this study, the parasitism of C. ohridella was examined in natural and artificial horse‐chestnut stands in South‐eastern Europe. The size and composition of the parasitoid complex of natural stands was comparable with that known from artificial stands. No specialist parasitoid species were detected in any of the samples. The only parasitoid species found exclusively in natural stands during this investigation were the braconid Colastes braconius and the eulophid Chrysocharis phryne. Both are known to be polyphagous on many leafminers in Europe and C. braconius has been often recorded from C. ohridella in artificial stands in previous studies. Parasitism levels varied within the same range (3.6–21.0%) throughout the entire region, in both types of habitats. 3 These results suggest that the parasitoid complex of C. ohridella in natural and planted horse‐chestnut stands in Europe is typically that of an exotic, recently introduced leafminer. Consequently, C. ohridella is probably not indigenous in the Balkans, but has been introduced to the area of its discovery from another continent.     

Pea leafminer Liriomyza huidobrensis (Diptera: Agromyzidae) uses vibrational duets for efficient sexual communication

The pea leafminer (Liriomyza huidobrensis) is a notorious pest of vegetables and ornamental plants worldwide. Despite a large number of studies on its biology and ecology, the courtship behavior and sexual communication of this species remain unclear. Here, we studied vibrational communication in the sexual interaction of the pea leafminer. On host plant leaves, females and males behaviorally displayed the bobbing-quivering alternation, which finally led to copulation. Moreover, records of laser vibrometry revealed threesignal duets underlying the behavioral alternation. Sexually mature males spontaneously emitted calls (MCs) to initiate the duets. The females rapidly responded to MCs by emitting replies (FRs) that are longer in duration. The FRs further triggered male replies (MRs) in their search for potential partners. Leafminer-produced vibrational signals convey efficient information to partners and generate pair formation on stretched substrates, such as plant leaves and nylon mesh, but cannot elicit responses on dense substrates, such as glass and plastic. Vibrational playbacks of both MCs and FRs can elicit replies in females and males, respectively. This study completely characterizes substrate-borne vibrational duets in a dipteran insect. The discovery of vibrational sex signals in the pea leafminer provides new insights for the development of novel approaches to control the pest and its relative species.     

First record of a walnut shield bearer Coptodisca (Lepidoptera: Heliozelidae) in Europe

A leafminer of the Nearctic genus Coptodisca Walsingham (Lepidoptera: Heliozelidae), a species of potential economic interest, is reported for the first time from Europe, infesting the black (Juglans nigra L.) and the common walnut (Juglans regia L.). Mines were collected since September of 2010 in several sites of two Italian regions (Campania and Lazio). The species is rather similar to Coptodisca juglandella (Chambers), the only Coptodisca known to attack walnuts, but at present, an unambiguous identification cannot be provided because of the unsatisfactory characterization of this leafminer and congeneric species. Three generations were recorded per year and leafminers overwinter as mature larvae. The first adults emerged in May–June while mature larvae of the last generation started the overwintering in September. During the last generation of the year, infestation levels of leaves were 100% in all sampled localities. Several species of parasitoids were reared from infested mines, with specimens belonging to the genus Chrysocharis (Hymenoptera: Eulophidae) being the most frequent parasitoids.     

Evaluation of the community of native eulophid parasitoids on Cameraria ohridella Deschka and Dimic in urban areas

The parasitoid complex associated with the exotic leafminer Cameraria ohridella Deschka and Dimic (Lepidoptera: Gracillariidae), which attacks horse chestnut (Aesculus hippocastanum L.), was studied in the urban environment of Turin (northern Italy). The studies were carried out over 5 yr after the first detection of the pest in our region in 1999. To evaluate parasitism, 438,029 leaf mines were examined over the 5-yr period, of which 29,033 were found to be parasitized (6.6%). Also, ornamental broadleaf trees attacked by other native gracillariid leafminers and located in the proximity of the target horse chestnut trees were sampled. A total of 11 parasitoid species (Hymenoptera: Eulophidae) were recorded on C. ohridella, and the most common species were Minotetrastichus frontalis (Nees), Closterocerus trifasciatus Westwood, and Pnigalio agraules (Walker). The first species accounted for >77.5% of all parasitoids collected. Cirrospilus talitzkii Boucek was found for the first time in 2005. The high population level of the pest and the low parasitism rate show that the parasitoid complex is currently inadequate to contain C. ohridella populations effectively. The most frequent parasitoids of the moth were also found on the most common broadleaf trees in the studied area, showing how native leafminer parasitoid species are able to switch to other hosts. These results show that both native and broadleaf plants species may potentially provide an important reservoir of parasitic wasps to help protect a simple biotope, such as the urban environment, from pests.     

Laboratory rearing and biological parameters of the eulophid Pnigalio agraules,a parasitoid of Cameraria ohridella

Predators as well as parasitoids native to Europe accept the exotic horse chestnut leafminer, Cameraria ohridella Deschka and Dimi? 1986 (Lepidoptera: Gracillariidae), either as prey or as host. However, the influence of these antagonists on the populations of the pest insect is so far very low. Therefore, efforts to develop an integrated pest management system against C. ohridella should include methods which foster the natural enemy complex. In the present study we developed a laboratory rearing method and investigated several biological parameters of Pnigalio agraules (Walker 1839) (Hymenoptera: Eulophidae), a dominant species in the horse chestnut leafminer’s parasitoid complex in many European regions. This native parasitoid satisfies three basic requirements for successful use as a biocontrol agent with regard to C. ohridella. The parasitoid’s fecundity, longevity and the speed of juvenile development by far exceeds that of the leafminer, enabling the parasitoid population to increase faster than that of the host. Furthermore, our results show that the impact of this species on C. ohridella populations has been previously underestimated, because non‐reproductive killing (i.e. host‐feeding and host‐stinging) of the hosts, resulting in considerable larval mortality of the leafminer, has not been quantitatively assessed. However, naturally occurring parasitoid populations have negligible impact on C. ohridella populations. Further studies, including experimental releases of P. agraules, are necessary to understand the constraints limiting the parasitoid’s performance in the field and to assess the potential benefits of releases for the control of C. ohridella.     

Cross‐resistance of chrysanthemum to western flower thrips,celery leafminer,and two‐spotted spider mite

Chrysanthemum [Chrysanthemum × morifolium Ramat. (Asteraceae)] is one of the economically most important greenhouse ornamentals worldwide. A major constraint in chrysanthemum production is adequate pest management, requiring the use of different tactics, such as improving host plant resistance, in the framework of an integrated pest management (IPM) approach. In this study, we investigated cross‐resistance of chrysanthemum to its three major pests: western flower thrips [Frankliniella occidentalis (Pergande) (Thysanoptera: Thripidae)], celery leafminer [Liriomyza trifolii (Burgess) (Diptera: Agromyzidae)], and two‐spotted spider mite [Tetranychus urticae Koch (Acari: Tetranychidae)]. We quantified resistance to each pest by performing greenhouse bioassays with a broad range of chrysanthemum types from commercial germplasm provided by Dutch breeding companies. Considerable variation was detected among the chrysanthemum cultivars in thrips silver damage and growth damage, leafminer damage, measured as number of mines and pupae, and spider mite numbers and damage. We observed significant positive correlations between thrips damage (both silver and growth damage) vs. leafminer numbers (both mines and pupae), and between leafminer numbers (both mines and pupae) vs. spider mite numbers. Our results indicate an overlap in resistance to all three herbivores. The important implications of this result for chrysanthemum breeding are discussed.     

Natural mortality factors of Leucoptera coffeella (Lepidoptera: Lyonetiidae) on Coffea arabica

The natural mortality of the coffee leafminer, Leucoptera coffeella (Guérin-Méneville) was investigated in three strata of coffee plant canopy for three seasons through construction and analysis of ecological life tables. Mortality of the leafminer was similar on all thirds of the canopy. Total mortality of immature stages was 95%, with 38.5, 43.8 and 12.7% occurring during egg, larval, and pupal stages. Rainfall killed 39.3% of eggs and larvae, and together with egg inviability (16.3%) and Vespidae (11.3%), were the highest mortality factors. Six wasp parasitoids caused 8% of larval mortality. Egg and larva were the critical stages. Variation in mortality was primarily associated with egg inviability, rainfall, and parasitism by Horismenus sp. (Hymenoptera: Eulophidae). Physiological disturbances during molting and metamorphosis also contributed for fluctuations in mortality of the leafminer. Tactics of integrated pest management to enhance natural mortality of the leafminer while conserving or augmenting the action of natural enemies are discussed.