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.     

外来斑潜蝇的优势寄生蜂——冈崎姬小蜂生物学特性及应用研究进展

冈崎姬小蜂是我国多种潜叶蝇的优势寄生蜂。鉴于蔬菜潜叶蝇尤其是入侵斑潜蝇在我国的肆意扩散和危害,本文总结了冈崎姬小蜂的分类地位、分布和田间发生、寄主种类、取食和繁殖、性比等生物生态学特性和田间应用进展。冈崎姬小蜂是一种抑性、卵育型的幼虫内寄生蜂,通过产卵寄生和取食2种方式致死寄主;雌蜂对寄主的偏好具有“寄主大小依赖型性别分配”现象;寄生蜂具有广泛的温度适应性和极强的控害潜力,尤其适用于控制入侵我国的相对耐高温的美洲斑潜蝇和三叶草斑潜蝇。未来将以更高效利用该寄生蜂为目标,主要集中于：（1）生物生态学和环境适应性的全面深入研究;（2）雌蜂寄主取食行为特性及生理机制的研究;（3）因地制宜的规模化饲养技术和释放应用技术研究;（4）与其他潜叶蝇寄生蜂的协同控害及竞争共存机制研究。     

Diversity, function and stability in parasitoid communities

The parasitoid assemblages associated with grass-feeding chalcid wasps in Great Britain were used to examine the relationships between diversity (species richness), community function (total parasitism rate) and stability (variability in parasitism rate over time). Species-rich communities did not generate higher parasitism rates than species-poor communities, nor was temporal variation of parasitism rates related to parasitoid community richness. The mechanisms underlying hypotheses linking species richness and community function and stability are discussed in the light of these results. Because all parasitoid species represent a single functional group, a lack of complementarity in the ways they use their resources may explain why diversity is not linked to function or community stability. A second likely reason is that these parasitoid communities are under bottom-up control, thus exerting little or no influence on total system function and variability. This is likely to be common in parasitoid communities.     

Geographic and between-generation variation in the parasitoid communities associated with an invading gallwasp,Andricus quercuscalicis (Hymenoptera: Cynipidae)

The knopper gallwasp Andricus quercuscalicis Burgsdorf 1783 (Hymenoptera: Cynipidae) has invaded western and northern Europe from southern and eastern Europe over the last 400 years. A. quercuscalicis has two alternating generations, which differ in phenology, structure, and host oak species. This study describes geographic variation in the community in the tiny catkin galls of the sexual generation on Turkey oak, Quercus cerris, and compares the patterns obtained with those in the community attacking the alternate agamic generation. As predicted from considerations of parasitoid recruitment to the communities of invading phytophagous insects (Cornell and Hawkins 1993), in its native range the sexual generation shows (1) higher parasitoid community species richness, (2) higher total mortality due to parasitoid attack and (3) a higher ratio of specialist to generalist parasitoid species than is evident in the invaded range. Counter to predictions, there is no indication that parasitoid community richness in the invaded range has increased with time since the arrival of the new host. Higher host mortality in the native range is due principally to a single specialist, Aulogymnus obscuripes Mayr 1877 (Hymenoptera: Eulophidae), and is not distributed evenly among parasitoid species which attack the gall-former only in this area. This contrasts with the community in Britain, where three principal generalist parasitoids cause approximately equal mortalities. The agamic gall contains a taxonomically and structurally diverse guild of parasitoid and inquiline species, associated with the changing resource provided by a large, long-lived, complex gall. In contrast, the sexual community includes a taxonomically and structurally narrow guild, associated with a resource which is structurally simple, small in size and short-lived. No parasitoid species attacks the gall-former in both generations. Surprisingly, in spite of these differences in the nature of the gall resource in the two generations, over their entire range (native and invaded) the parasitoid guilds of the two are equally species rich.     

Redescription of Closterocerus cruy (Girault) (Hymenoptera: Eulophidae: Entedoninae), with new geographical and host records

Abstract  Closterocerus cruy (Girault) is newly recorded, both from New Zealand and as a parasitoid of lepidopteran leafminers. The species is redescribed from the Girault type and from New Zealand material. Diagnostic character states are provided for the separation of C. cruy from the closely related Australian agromyzid parasitoid C. mirabilis Edwards and LaSalle.     

Community ecology of metazoan parasites of the anchovy Anchoa tricolor (Osteichthyes: Engraulidae) from the coastal zone of the State of Rio de Janeiro, Brazil.

Between October 2001 and March 2002, 103 specimens of A. tricolor from Angra dos Reis (23 degrees 01' S, 44 degrees 19' W), in the coastal zone of the State of Rio de Janeiro, Brazil, were analyzed in order to study their metazoan parasite infracommunities. Ten species of metazoan parasites were collected: 4 digeneans, 1 cestode, 1 acantocephalan, 2 nematodes, 1 copepod, and 1 hirudinean; 77.7% of the fishes were parasitized by one or more metazoan, with a mean of 3.5 +/- 6.2 parasite/fish. Digenean was the most dominant with 4 species that accounted for 53.2% of the total parasites collected; Ergasilus sp. was the most abundant species. Abundance and prevalence of Parahemiurus merus (Linton, 1910) were positively correlated with the total length of host. Relationships between total body length of fish and both total parasite abundance and mean parasite species richness were observed. Mean parasite diversity of species was correlated to host's total length, with significant differences found between male and female fishes. Two pairs of larval species showed significant positive association and covariation. The metazoan parasite infracommunities of A. tricolor presented dominance of larval endoparasites; correlation of parasite abundance, diversity, and species richness with host total length; and low number of parasite interspecific relationships. The parasite community of A. tricolor showed some similarities with the parasite community of another South American Atlantic engraulid.     

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.     

Edge effects on insect–plant food webs: assessing the influence of geographical orientation and microclimatic conditions

1. Geographical orientation affects the magnitude of microclimatic edge effects, mainly by altering the exposure to solar radiation. There is considerable evidence available concerning the influence of microclimatic edge effects on organisms at the population level, although little is known about effects at the community level, and variations with edge geographical orientation have barely been addressed. The present study investigated the influence of microclimatic conditions at the forest interior and edges with different geographical orientation on plant–leafminer–parasitoid food webs in the fragmented Chaco Serrano forest located in Central Argentina. 2. Microclimatic variables were measured in three locations: interior, north, and south-facing edges of six forest fragments. In each location, plant resource availability was assessed and mined leaves were collected to investigate plant–leafminer–parasitoid communities by rearing leafminer and parasitoid adults. 3. The results obtained revealed changes in the specific composition of plant, leafminer and parasitoid communities among locations, although they failed to detect location influences on most variables linked to abundance (except for leafminers), richness or food web structure. Leafminer abundance was negatively related to temperature, whereas their richness was related, also negatively, to moisture. Instead, parasitoids were favoured by higher temperatures and light levels, stronger winds, and lower humidity. 4. The results of the present study indicate that the effects of microclimatic conditions may depend on the trophic level considered. Moreover, the findings suggest that possible trade-offs between direct and indirect effects (such as plant quality, floral resources, etc.) of microclimatic conditions could influence plant and insect responses to location.     

Intraspecific size variation in polyphagous parasitoids (Hym.: Parasitica), of leaf miners and its relation to host size

Wing and body lengths of polyphagous parasitoids of leafminers are analyzed in order to understand the relationship between host and parasitoid sizes. A distinct positive relationship was observed: within each parasitoid species, smaller individuals were reared from smaller hosts. There was no difference between idio-and koinobionts in sexual dimorphism, with females being significantly larger than males in both groups.     

Effects of parasitoid community structure upon the population dynamics of the honeysuckle leafminer,Chromatomyia suikazurae (Diptera: Agromyzidae)

Population dynamics of a leafminer,Chromatomyia suikazurae (Agromyzidae, Diptera) and its parasitoid community were studied for ten years at seven natural populations along an altitudinal gradient in Japan. This species which mines leaves of a forest shrub,Lonicera gracilipes (Caprifoliaceae), was attacked by 25 hymenopterous parasitoid species. Annually, the parasitoid community structure varied less within a population than among populations. The seven parasitoid communities were clustered into three groups corresponding to the altitudinal gradient: (a) lowland communities dominated by late-attacking, generalist pupal idiobiont eulophids and with highest species diversity, (b) hillside communities dominated by an early-attacking, specialist larval-pupal koinobiont braconid and (c) highland communities dominated by an early-attacking, generalist larval idiobiont eulophid. Annual changes of the host larval densities among the local populations were largely synchronous rather than cyclic. Among these populations, host density levels and mortality patterns greatly varied. By analyzing these inter-populational differences of host mortality patterns, the following conclusions were drawn: (1) The host mortality patterns were determined by the host utilization patterns of the locally dominant species. (2) The host pupal mortality but not larval mortality was related to species diversity but not to species richness itself of each parasitoid community. (3) Density dependence was detected only in pupal mortality at a lowland population dominated by late-attacking pupal parasitoids. These results suggest that interspecific interactions of parasitoids add additive effects to host population dynamics dissimilarly among local populations with different parasitoid communities.     

Parasitic hymenoptera fauna on agromyzidae (diptera) colonizing weeds in ecological compensation areas in northern Italian agroecosystems

Parasitoids (Hymenoptera) associated with agromyzid leafminers (Diptera: Agromyzidae) were studied in three rural farms located in northern Italy. The parasitoids were reared from mined foliage of weeds growing in field margins. We reared 998 Hymenoptera specimens, representing five families, 23 genera, and 53 species, from leafminers infesting weeds. Eulophidae was the most abundant family (67.64%), followed by Braconidae (28.86%), Eucoilinae (1.40%), Tetracampidae (1.40%), and Pteromalidae (0.7%). Braconids was the most species rich family, accounting for 28 species; eulophids were represented by 19 species, pteromalids by four species, and eucoilins and tetracampids by one species each. The dominant parasitoid was the eulophid Pediobius metallicus (Nees), representing 18.17% of the total, followed by Diglyphus isaea (Walker) (12.73%), and Neochrysocharis formosa (Westwood) (10.82%). The most abundant braconid parasitoid was Dacnusa maculipes Thomson (9.62%). More than 80% of parasitoids were recovered from 10 plant species: Cirsium arvense (L.) Scopoli, Plantago lanceolata L., Sonchus asper (L.) Hill, Papaver rhoeas L., Picris echioides L., Lactuca serriola L., Myagrum perfoliatum L., Ranunculus velutinus Tenore, Arctium lappa L., and Medicago sativa L. The retention and the management of wild plants within field margins can be crucial tools to enhance the populations of biological control agents of agromyzids and to conserve rare parasitic wasp species.     

The occurrence of leafminers and their parasitoids on vegetables and weeds in Hangzhou area, Southeast China

Leafminers are the mostimportant insect pests of vegetables inHangzhou area, Zhejiang province, SoutheastChina as well as in the rest of China, but havenever been investigated for speciescomposition, relative abundance and theirassociated parasitoids. A survey was conductedin four localities of Hangzhou area during 1998–2000to determine the distribution ofleafminers and their parasitoids on vegetablecrops and weeds in the field. Over 20,000leafminers were collected and reared. Theleafminer species found were Liriomyzasativae, L. chinensis and Chromatomyia horticola, with C.horticola and L. sativae the mostabundant in spring and autumn, respectively.About 11,000 parasitoid adults were reared fromthe leafminers collected. In total 14hymenopteran species of four families (onebraconid, 10 eulophids, one megaspilid and onepteromalid) were found. Eleven species werereared from C. horticola, nine from L. sativae and seven from L. chinensis.Opius caricivorae and Chrysocharispentheus were the dominant species. Sevenspecies were recorded from L. chinensisfor the first time. Eleven parasitoid specieswere reared from leafminers on two weeds (Veronica undulata and Sonchusoleraceus). Parasitoids caused 48.5–68.8%parasitism on leafminers on crops and 83.7%parasitism on leafminers on weeds in the latergrowing season of vegetable crops. The resultssuggest that leafminer populations areregulated to a certain extent by their naturalenemies in the field.     

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.

     

Host diversity and environmental variables as determinants of the species richness of the parasitoids of leaf‐cutting ants

Aim Because of the obligatory relationship between endoparasitoids and their hosts, we presume that hosts exert strong selection pressure on parasitoids. One prediction is a positive relationship between host diversity and parasitoid richness. This relationship could be the product of resource availability which could lead to more opportunities for speciation, or could represent shared responses to the environment by both groups. Location Argentina and Paraguay. Methods We sampled a 1800‐km transect to test for a correlation between the richness of leaf‐cutting ant hosts and their phorid parasitoids. Regression models were used to assess if host and environmental variables could explain phorid species richness at nest, hectare and locality spatial scales. We used canonical correspondence analysis (CCA) to explore if there were similar responses of phorid species to particular host and environmental variables at different spatial scales, and partial CCA to separate the relative importance of both groups of variables. Results Phorid richness was positively correlated with host richness. Host richness/abundance accounted for 20–53% of the variation in parasitoid richness at the hectare and locality scales of analysis, with most of the variation accounted for by ant abundance. We were not able to assess the prediction at the nest scale as only one phorid species was found at most nests. Climatic variables did not explain phorid species richness once host variables were in the models. Partial CCA showed that host‐related variables accounted for most of the variance associated with phorid species ordination at the nest and hectare scales, but not at the largest grain, the locality, where climatic variables were more important. However, most phorid species did not show particular positions along the climatic gradient. Main conclusions The association between parasitoid richness and host richness and abundance, and the overall weak associations with environmental variables, suggest that these host variables are key factors influencing parasitoid speciation.     

Effects of landscape structure and habitat type on a plant-herbivore-parasitoid community

The effects of local habitat and large-scale landscape factors on species diversity and species interactions were studied using the insect community in stems of the creeping thistle Cirsium arvense . Thistle abundance was higher in fallows than in crop fields and field margins, with fallows providing 67% of thistle abundance within 15 study areas on a landscape scale. Species richness of the herbivores was positively related with thistle abundance, parasitoid species richness was influenced by habitat type and was positively correlated with herbivore species richness. The abundance of herbivores and parasitoids was affected by local factors such as habitat type and host abundance, but also by landscape factors such as the percentage of non-crop area and the isolation of habitats. The infestation rate caused by the agromyzid Melanagromyza aeneoventris was positively related to percent non-crop area, whereas the parasitism rate of this fly increased with increasing habitat diversity on the landscape scale. For these two interactions and for total herbivore abundance, a scale-dependency of the landscape effects was found. The results emphasize that biological diversity and ecological functions within a plant-insect community are not only affected by local habitat factors but also by large-scale landscape characteristics. Hence, to improve future agri-environmental schemes for biodiversity conservation and biological control large-scale landscape effects and their scale-dependency should be considered.     

潜蝇姬小蜂属寄生蜂对潜叶蝇的控害特性及应用

潜蝇姬小蜂属Diglyphus寄生蜂是潜叶蝇类害虫的重要天敌。本文对其种类、 分布、 田间发生和优势度、 优势种的控害特性和生态适应性, 以及优势种的人工繁殖和田间应用等进行总结和展望, 以期为更好地应用该属寄生蜂防控我国潜叶蝇的研究和应用提供指导。迄今已鉴定该属寄生蜂36种, 全为抑性外寄生蜂, 其中针对豌豆潜蝇姬小蜂D. isaea、 贝氏潜蝇姬小蜂D. begini和中带潜蝇姬小蜂D. intermedius的研究较多。雌蜂不仅可通过繁殖型的寄生方式而且还可通过非繁殖型方式（取食寄主和产卵器插入直接杀死）致死寄主。雌蜂偏好寄生大个体寄主和取食相对较小个体的寄主, 且雌蜂对寄主幼虫的偏好具有“寄主大小依赖型性别分配”现象（host-size-dependent sex allocation）。从温度适应范围和控害潜力首推为豌豆潜蝇姬小蜂。优势种的规模饲养技术已经被研发并得到了较广泛的田间应用。未来研究可主要集中于： （1）加强优势种尤其是温度适应范围广或较为耐热的种群或地理品系的研究, 以增加对三叶草斑潜蝇Liriomyza trifolii和美洲斑潜蝇L. sativae的控效; （2）加强雌蜂寄主取食行为特性及其生理机制的研究, 以更高效利用; （3）因地制宜研发规模化饲养技术和释放技术; （4）加强潜蝇姬小蜂同其他潜叶蝇寄生蜂的协同控害及竞争共存机制的研究, 已提升对潜叶蝇的生防控制效果。     

Determinants of assemblage size for the parasitoids of Cecidomyiidae (Diptera)

Summary The parasitoids known to attack 191 phytophagous species of gall midges (Cecidomyiidae) were used to examine factors influencing parasitoid assemblage size. The number of parasitoid species a midge species supports was tested against nine variables describing geographical, biological and ecological attributes of hosts. The apparency of midge larvae was found to have the greatest influence on parasitoid assemblage size; highly visible species support more parasitoids than less visible ones. Pupation site and midge voltinism also significantly affect associated parasitoids, at least for highly apparent hosts. Biogeographic region, host-plant architecture and the plant parts infested were found to be of secondary importance. The surface texture of infested plant parts, the number of midge larvae occupying galls and the diversity of plant tissues infested have minimal apparent effects on parasitoid richness. Parasitoid assemblage size and total parasitism rates were also found to be positively correlated for 73 galling and nongalling midge species, and gallers typically suffer higher levels of parasitism than non-gallers. Using these data to test the enemy hypothesis, which proposes that the galling habit has evolved to escape attack from parasitoids, we conclude that parasitoid pressure cannot account for the presence of galls in the Cecidomyiidae.     

Occurrence and phenology of parasitic Chalcidoidea on the horse chestnut leafminer,Cameraria ohridella Deschka & Dimic (Lep., Gracillariidae)

Abstract: The parasitoid spectrum of the horse chestnut leafminer, Cameraria ohridella, was examined for its adaptation to a newly introduced host. A total of 15 parasitic species belonging to the suprafamilies Ichneumonoidea and Chalcidoidea was reared, all of them are polyphagous and common on other leafminers in Europe. The abundance of the moth and its natural enemies were studied during one entire vegetation period. The phenology of some of its major parasitic species was investigated and compared with the occurrence of the leafminer with special regard to its possible use in biological pest control.     

Urbanization Level and Woodland Size Are Major Drivers of Woodpecker Species Richness and Abundance

Urbanization is a process globally responsible for loss of biodiversity and for biological homogenization. Urbanization may have a direct negative impact on species behaviour and indirect effects on species populations through alterations of their habitats, for example patch size and habitat quality. Woodpeckers are species potentially susceptible to urbanization. These birds are mostly forest specialists and the development of urban areas in former forests may be an important factor influencing their richness and abundance, but documented examples are rare. In this study we investigated how woodpeckers responded to changes in forest habitats as a consequence of urbanization, namely size and isolation of habitat patches, and other within-patch characteristics. We selected 42 woodland patches in a gradient from a semi-natural rural landscape to the city centre of Poznań (Western Poland) in spring 2010. Both species richness and abundance of woodpeckers correlated positively to woodland patch area and negatively to increasing urbanization. Abundance of woodpeckers was also positively correlated with shrub cover and percentage of deciduous tree species. Furthermore, species richness and abundance of woodpeckers were highest at moderate values of canopy openness. Ordination analyses confirmed that urbanization level and woodland patch area were variables contributing most to species abundance in the woodpecker community. Similar results were obtained in presence-absence models for particular species. Thus, to sustain woodpecker species within cities it is important to keep woodland patches large, multi-layered and rich in deciduous tree species.