Plant hosts and parasitoid associations of leaf mining flies (Diptera: Agromyzidae) in the Canberra region of Australia

Abstract  Many leaf mining flies (Diptera: Agromyzidae) are important economic pests of agricultural crops and ornamental plants, and species-rich hymenopteran parasitoid complexes are important in their control. Australian agromyzids are poorly studied, and little is known about their host plants, ecology or natural enemies. We surveyed native and naturalised species of leaf mining flies in Tallaganda National Park, New South Wales and the Australian Capital Territory. Malaise and emergence trapping in Tallaganda yielded 70 agromyzid specimens from six species in four genera: Cerodontha Rondani, Liriomyza Mik, Phytoliriomyza Hendel and Phytomyza Fallen. Of the six species collected, three are Australasian species, two are naturalised species introduced from Europe and one could not be determined to species. The Australian Cerodontha ( Cerodontha ) milleri Spencer represented most of the individuals caught in both Malaise and emergence traps. A total of 163 agromyzid and 98 parasitic wasp specimens were reared from plant samples with agromyzid mines in the Canberra region. Most agromyzids and parasitoids were reared from the weed Sonchus oleraceus L. (Asteraceae). All the agromyzids reared belonged to two introduced species of the genera Phytomyza and Chromatomyia Hardy. The biodiversity of parasitic wasps reared was high with 14 species from seven genera and three families. Hemiptarsenus varicornis (Girault) (Eulophidae), a widespread Old World agromyzid parasitoid, was the most numerous parasitoid reared in our survey.     

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.     

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。     

Structure,organization, and response of a species-rich parasitoid community to host leafminer population dynamics

The parasitoid community dynamics of an agromyzid honeysuckle leafminer, Chromatomyia suikazurae (Agromyzidae, Diptera) were studied between 1981 and 1990 in a natural forest in Kyoto, Japan. The parasitoid fauna composed three koinobionts (all larval-pupal solitary parasitoids) and 22 idiodiont species (11 larval solitary, nine pupal solitary and one pupal gregarious). The parasitoid community was dominated by early-attacking oligophagous braconid koinobionts at early periods, but was gradually displaced by late-attacking polyphagous eulophid idiobionts. Accordingly, the diversity index of the parasitoid community peaked at an intermediate point in the intra-generational succession. The succeeding attack-in-waves by the late-attacking idiobionts greatly reduced not only the survival rates of early-attacking parasitoid larvae but also the survival rates of hosts. The density-dependence observed in the host pupal mortality was thought to result from density-dependent host-switching by a keystone polyphagous pupal idiobiont parasitoid, Chrysocharis pubens, whereas high host pupal mortality was potentially attained by an early-attacking koinobiont braconid. Supposed aggregation of polyphagous parasitoids at high host density resulted in intense within-host competition and in an increase of host-feeding attack, both of which contributed to low emergence rates of parasitoids at high host densities. Parasitoid emergence rates were also reduced at low host densities, probably by inter- and intra-specific hyperparasitism among oligophagous parasitoids for limited hosts. The regulation effects of the species-rich parasitoid community upon the host population dynamics are thought to derive from succeeding attack-in-waves by polyphagous late-attacking idiobionts, especially by the keystone species.     

甘肃白银地区潜叶蝇及其寄生蜂的组成和发生调查

【目的】明确严重危害甘肃省白银地区作物的潜叶蝇及天敌寄生蜂的多样性和种群组成。【方法】采用五点取样法田间调查潜叶蝇危害的蔬菜、花卉、杂草并采样,室内对潜叶蝇及其寄生蜂进行形态鉴定和分子鉴定。【结果】在2016-2020年在甘肃白银地区采集的样本中发现6种常见潜叶蝇,包括本地种豌豆彩潜蝇Chromatomyia horticola和葱斑潜蝇Liriomyza chinensis,入侵种南美斑潜蝇L. huidobrensis、三叶草斑潜蝇L. trifolii、番茄斑潜蝇L. bryoniae和美洲斑潜蝇L. sativae。其中,豌豆彩潜蝇为优势种,可危害多科寄主植物;葱斑潜蝇仅在大葱上发现危害;南美斑潜蝇在温室内的蔬菜上发生严重;三叶草斑潜蝇首次在白银市发现;番茄斑潜蝇与美洲斑潜蝇的危害相对较轻。此外,调查的潜叶蝇寄生蜂包括3科12属24种,优势种包括豌豆潜蝇姬小蜂Diglyphus isaea、芙新姬小蜂Neochrysocharis formosa、万氏潜蝇姬小蜂D. wani和西伯利亚离颚茧蜂Dacnusa sibirica。【结论】一方面白银市应加强监测和预警入侵性潜叶蝇的发生,另一方面白银市潜叶蝇的天敌寄生蜂种类丰富,对田间主要危害的潜叶蝇可能发挥着重要的自然防控作用,建议保护利用本地寄生蜂天敌。     

Diglyphus isaea (Hymenoptera: Eulophidae): a probable complex of cryptic species that forms an important biological control agent of agromyzid leaf miners

Diglyphus isaea ( Walker 1838 ) (Hymenoptera: Eulophidae) is a primary parasitoid of agromyzid leaf miners (Diptera: Agromyzidae) and has been commercialized as a biological control agent. Diglyphus isaea occurs throughout much of the world and different populations are morphologically identical. Using nuclear ribosomal internal transcribed spacer 1 (ITS1) sequences, we examined variation among Chinese populations of D. isaea . Phylogenetic analyses combined with an analysis of sympatry indicated that D. isaea consists of at least four species in mainland China. The results imply that ITS1 is an efficient marker for identifying congeneric species of parasitic waSPS, and that cryptic species could be common in temperate and subtropical regions.     

Native saltbush (Rhagodia spp.; Chenopodiaceae) as a potential reservoir for agromyzid leafminer parasitoids on horticultural farms

Australia has to date been spared the introduction of highly polyphagous invasive pest agromyzid leafminers; however, their arrival and spread should be considered imminent. To develop a pre-emptive control strategy to deal with exotic leafminer outbreaks the first step is to identify Australian leafmining flies, their plant hosts and their parasitoids to gain an understanding of their population dynamics. Native vegetation may be providing resources for beneficial parasitic wasps plus access to alternative hosts and refuge from disturbance. Here, two Australian endemic saltbushes ( Rhagodia candolleana and R. parabolica , Caryophyllales: Chenopodiaceae) have been investigated for their potential to act as reservoirs for endemic agromyzid hosts and their key parasitoids. Mined leaves of the two Rhagodia species were sampled on two commercial horticultural properties in the Virginia horticulture area on the Northern Adelaide Plains between September 2007 and April 2008. Leaf mines on both Rhagodia species were caused by an endemic leafminer species, putatively Phytoliriomyza praecellens Spencer (Diptera: Agromyzidae). Ten species of parasitoids (all Hymenoptera) emerged from R. candolleana mines and seven different species from R. parabolica mines, mainly from the family Eulophidae and with some Pteromalidae and Braconidae. Trigonogastrella Girault sp. (Pteromalidae), Zagrammosoma latilineatum Ubaidillah and Hemiptarsenus varicornis Girault (both Eulophidae) were the most abundant species on R. candolleana , whereas two Opius Wesmael spp. (Braconidae) were the most abundant species on R. parabolica . Findings from this survey suggest an opportunity to plant purpose-designed refuges that could play a role in conservation biological control as part of an Integrated Pest Management strategy developed prior to incursion of pest leafminers such as Liriomyza species.     

Community dynamics of leafminers (Diptera: Agromyzidae) and their parasitoids (Hymenoptera) in a natural habitat from Central Argentina

We examined the temporal community dynamics of leafminers (Agromyzidae: Diptera) and their mainly polyphagous parasitoids in a natural habitat from Central Argentina. Changes in community composition are shown by changes in dominant species and similarity coefficients. Abundance of leafminers was highest in winter, and was not related to their species richness. Abundance and species richness were correlated in the parasitoid community. Temporal variation in parasitoid diversity and abundance were positively correlated with species number and density in the host community. Apparent parasitism was greater when parasitoid species richness was high and parasitoid community dominance was low.     

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.     

New host-plant records for neotropical agromyzids (Diptera: Agromyzidae) from Asteraceae flower heads

Agromyzidae is a large and cosmopolitan fly family with approximately 2,500 known species. Here we present 22 new records of agromyzid-host plant associations. Plants were sampled from 2002 to 2005 in S?o Paulo state, Brazil. A total of eight agromyzid species were reared from 18 Asteraceae host species. The genus Melanagromyza Hendel was the commonest. This is the first detailed study reporting associations between non-leafmining Agromyzidae and their host plants in Brazil.     

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。     

Taxonomic Composition of Hymenopteran Parasitoid Assemblages from Agromyzid Leaf-miners Sampled in Central Argentina

This paper deals with the taxonomic composition of parasitoid assemblages reared from agromyzid leaf-miners of 28 species collected in Cordoba, Central Argentina. The relative representation of each parasitic family was estimated, taking into consideration their species richness and abundance. Sample size affected the total species richness in each complex, and also that of Eulophidae and Pteromalidae, indicating that not only size but also taxonomic structure of parasitic assemblages are influenced by sampling effort. 3 superfamilies and 5 families of Hymenoptera Parasitica were represented, with Eulophidae (Chalcidoidea) specifically and numerically dominant both in the total community of parasitoids and in the parasitic assemblages of individual hosts. Chrysonotomyia, Chrysocharis (Eulophidae), Halticoptera (Pteromalidae) and Phaedrotoma (Braconidae) were the most relevant genera in terms of number of species and number of specimens reared. The parasitoids obtained from samples taken in agroecosystems differed from the global spectrum of taxa mainly by showing a numerical dominance of Braconidae, despite a noticeable specific impoverishment of this family.     

Efficacy of using DNA barcoding to identify parasitoid wasps of the melon-cotton aphid (<Emphasis Type="Italic">Aphis gossypii</Emphasis>) in watermelon cropping system

Parasitoid wasps have received a great deal of attention in the biological control of melon-cotton aphid (Aphis gossypii Glover). The species of parasitoids are often difficult to identify because of their small body size and profound diversity. DNA barcoding offers scientists who are not expert taxonomists a powerful tool to render their field studies more accurate. Using DNA barcodes to identify aphid parasitoid wasps in specific cropping systems may provide valuable information for biological control. Here, we report the use of DNA barcoding to confirm the morphological identification of 14 species (belonging to 13 genera of 7 families) of parasitoid wasps from two-year field samples in a watermelon cropping system. We generated DNA sequences from the mitochondrial COI gene and the nuclear D2 region of 28S rDNA to assess the genetic variation within and between parasitoid species. Automatic Barcode Gap Discovery (ABGD) supported the presence of 14 genetically distinct groups in the dataset. Among the COI sequences, we found no overlap between the maximum K2P distance within species (0.49%) and minimum distance between species (6.85%). The 28S sequences also showed greater interspecific distance than intraspecific distance. DNA barcoding confirmed the morphological identification. However, inconsistency and ambiguity of taxonomic information available in the online databases has limited the successful use of DNA barcoding. Only five species matched those in the BOLD and GenBank. Four species did not match the entries in GenBank and five species showed ambiguous results in BOLD due to confusing nomenclature. We suggested that species identification based on DNA barcodes should be performed using both COI and other genes. Nonetheless, we demonstrate the potential of the DNA barcoding approach to confirm field identifications and to provide a foundation for studies aimed at improving the understanding of the biocontrol services provided by parasitoids in the melon ecosystem.     

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.     

三叶草斑潜蝇的寄生蜂及其应用

本文就三叶草斑潜蝇Liriomyza trifolii(Burgess)的寄生蜂种类、生物学特性、寄主选择性和农药对寄生蜂的影响及寄生蜂在生物防治中的应用等方面的研究进展进行了综述。     

Histories of host shifts and cospeciation among free‐living parasitoids of Rhagoletis flies

Host shifts by specialist insects can lead to reproductive isolation between insect populations that use different hosts, promoting diversification. When both a phytophagous insect and its ancestrally associated parasitoid shift to the same novel host plant, they may cospeciate. However, because adult parasitoids are free living, they can also colonize novel host insects and diversify independent of their ancestral host insect. Although shifts of parasitoids to new insect hosts have been documented in ecological time, the long‐term importance of such shifts to parasitoid diversity has not been evaluated. We used a genus of flies with a history of speciation via host shifting (Rhagoletis [Diptera: Tephritidae]) and three associated hymenopteran parasitoid genera (Diachasma, Coptera and Utetes) to examine cophylogenetic relationships between parasitoids and their host insects. We inferred phylogenies of Rhagoletis, Diachasma, Coptera and Utetes and used distance‐based cophylogenetic methods (ParaFit and PACo) to assess congruence between fly and parasitoid trees. We used an event‐based method with a free‐living parasitoid cost model to reconstruct cophylogenetic histories of each parasitoid genus and Rhagoletis. We found that the current species diversity and host–parasitoid associations between the Rhagoletis flies and parasitoids are the primary result of ancient cospeciation events. Parasitoid shifts to ancestrally unrelated hosts primarily occur near the branch tips, suggesting that host shifts contribute to recent parasitoid species diversity but that these lineages may not persist over longer time periods. Our analyses also stress the importance of biologically informed cost models when investigating the coevolutionary histories of hosts and free‐living parasitoids.     

The response of aphid secondary parasitoids to different patch densities of their host

Aphids are attacked by a large guild of natural enemies including many primary parasitoids which mummify their hosts. These mummies are themselves attacked by a guild of mummy parasitoids which are potentially important in regulating primary parasitoids at densities below which they can exert biological control. The response of mummy parasitoids to mummy densities was investigated in an experiment in which mummy densities of the pea aphid (Acyrthosiphon pisum) attacked by the parasitoid Aphidius ervi were manipulated across host plant patches. Overall, the risk of parasitism was density independent, though with very high inter-patch variability which may allow probabilistic refuges from secondary parasitism. Six species of four genera of mummy parasitoids were recorded. Of the responses of the individual genera, Coruna were reared most frequently from patches of high host density while amongst patches from which Syrphophagus was reared parasitism was inversely density dependent.     

The colonization of native phytophagous insects in North America by exotic parasitoids

Classical biological control could have a major environmental cost if introduced natural enemies colonize and disrupt native systems. Although quantifying these impacts is difficult for systems already colonized by natural enemies, the a priori condition for such impacts can be evaluated based on the extent to which exotics have acquired native hosts. We use native host records for exotic parasitoids introduced into North America for biological control to document the number of exotic species that have been recorded from at least one native insect species. We also evaluate the ability of six biological and ecological variables to predict whether or not a parasitoid will move onto natives. Sixteen percent of 313 parasitoid species introduced against holometabolous pests are known from natives. Further, the likelihood that a parasitoid had colonized native hosts was largely unpredictable with respect to the independent variables. We conclude that given the quality of the data available either now or in the foreseeable future, coupled with inherent stochasticity in host shifts by parasitoids, there are no rules of thumb to assist biological control workers in evaluating if an introduced parasitoid will colonize native insect communities. Received: 2 July 1997 / Accepted: 3 August 1997