Functional response of a parasitoid Ganaspidium utilis （Hymenoptera： Eucoilidae） on the leafminer Liriomyza trifolii （Diptera： Agromyzidae）

Functional response of a solitary, larval-pupal endoparasitoid of Liriomyza leafminers, Ganaspidium utilis Beardsley, was estimated on Liriomyza trifolii Burgess at three temperatures （17℃, 25℃, 29℃ ） and host densities. A type Ⅱ random parasitoid equation （RPE） was used to estimate instantaneous search rate and handling time. The instantaneous search rate increased as temperature increased. All of the RPE regressions obtained for functional response of G. utilis at different temperatures were significant （P〈0.01）. The slope of RPE regression lines was lower across the temperatures. At 29±2℃, the maximum number of larvae parasitized was 7.8 per day. It decreased to 7.2 larvae parasitized at 25±2℃. At 17±2℃, no significant increment of parasitization was observed due to the host density increments. The estimated handling time was lowest at 17±2℃ and highest at 25 ± 2℃, respectively. The ability of G. utilis to find and parasitize L. trifolii over a wide range of temperatures makes them a good candidate for biological control of Liriomyza leafminers.     

入侵害虫南美斑潜蝇在西藏首次发现及其寄生蜂调查

【目的】南美斑潜蝇Liriomyza huidobrensis为我国重要的入侵害虫,我们于2017年7月首次发现其已扩散至西藏。本研究旨在了解西藏地区南美斑潜蝇的发生现状及其寄生蜂种类。【方法】2017和2018年6月上旬-9月中旬对西藏拉萨、林芝、山南和日喀则4个地区的南美斑潜蝇的发生危害及其寄生蜂组成进行了调查采样。【结果】调查的西藏这4个地区均发现南美斑潜蝇分布和危害;温室发生时间为6月上旬-9月中旬,而露地的发生高峰在7月下旬-8月上旬。其寄主植物有9科21种,主要有豆科(Fabaceae)、茄科(Solanaceae)和伞形科(Umbelliferae)等,主要嗜好经济作物有豇豆Vigna unguiculata, 扁豆Lablab purpureus , 莴笋Lactuca sativa var. angustana和白菜Brassica rapa ssp. pekinensis。其寄生蜂包括姬小蜂科(Eulophidae)、茧蜂科(Braconidae)和瘿蜂科(Cynipidae)共计3科9属16种,分别为双斑潜蝇姬小蜂Diglyphus bimaculatus, 豌豆潜蝇姬小蜂Diglyphus isaea, 小斑潜蝇姬小蜂Diglyphus minoeus, 万氏潜蝇姬小蜂Diglyphus wani, Diglyphus sp., 纵纹瑟姬小蜂Cirrospilus vittatus, Hemiptarsenus sp., 芙新姬小蜂Neochrysocharis formosa, 底比斯姬小蜂Chrysocharis pentheus, 普金姬小蜂Chrysocharis pubicornis, Chrysocharis sp., 潜蝇柄腹姬小蜂Pediobius metallicus, Pnigalio sp., 西伯利亚离颚茧蜂Dacnusa sibirica, Dacnusa sp.和Gronotoma sp.;其中优势种为西伯利亚离颚茧蜂、双斑潜蝇姬小蜂和豌豆潜蝇姬小蜂,其个体数量占比分别为总采集个体数的48.7%, 22.1%和12.9%。【结论】西藏南美斑潜蝇的寄生蜂种类丰富,发挥着重要的自然控制作用。建议进一步加强优势寄生蜂生物学、控害潜力评价以及利用研究,为南美斑潜蝇的绿色防控提供指导。     

南美斑潜蝇寄主选择性与植物次生化合物及叶毛的关系

采用非自由选择法,以刺伤孔（包括取食孔和产卵孔）数为指标,在室内测定了南美斑潜蝇对6科、16种、22个蔬菜品种的寄主选择性。结果表明,南美斑潜蝇对供试蔬菜品种的选择性存在显著差异,选择性大小依次为豆科>菊科>葫芦科>茄科>伞形科>十字花科; 菜豆和茼蒿是南美斑潜蝇最喜欢产卵和取食的寄主,而甘蓝和番茄是其选择性最差的寄主。化学分析和解剖镜观察表明,叶片单宁酸、黄酮含量及叶毛密度在供试品种间存在显著差异： 苦瓜的单宁酸含量最高,其次为豌豆、番茄和青椒,含量最低的为金丝搅瓜、茼蒿、菜豆和美国西芹; 黄酮含量最高的为甘蓝和番茄,其次为苦瓜,最低为茼蒿、菜豆和美国西芹; 南瓜和小白菜的叶毛密度最高,其次为西葫芦、金丝搅瓜和甘蓝,最低为青椒、莴苣和苦瓜。相关分析表明,南美斑潜蝇寄主选择性与叶片单宁酸和黄酮含量呈显著的负相关关系,相关系数分别为-0.4425（ P=0.0392）和 -0.5728（ P=0.0053）,而与叶毛密度相关关系不显著（ R＝-0.1807,P=0.4211）。说明黄酮和单宁酸对南美斑潜蝇产卵和取食具有明显的抑制作用。     

福建省发现加藤斑潜蝇（双翅目：潜蝇科Diptera：Agromyzidae）

1995年在调查潜叶蝇种类时，在菊花上发现加藤斑潜蝇，属中国大陆首次报道，本文对其形态特征进行描述。     

Avermectin类农药对美洲斑潜蝇的生物活性

Avermectin类农药对美洲斑潜蝇Liriomyzasattvae Blanchard生物活性的室内研究结果表明： 揭阳霉素对美洲斑潜蝇1、2、3龄幼虫的LC₅₀分别是1.54×10^-4 g/L、3.73×10^-4 g/L、1.99×10^-3g/L;害极灭对上述幼虫的LC₅₀分别是1.48×10^-4g/L、3.68x10^-4 g/L和1.97×10^-3 g/L。美洲斑潜蝇幼虫对Avermectln类农药以1龄最敏感,其中揭阳霉素对3龄幼虫的LC₅₀是1龄的12.9倍。揭阳霉素对雌成虫24 h、48 h的LC₅₀分别是3.12×10^-3 g/L和2.08×10^-3 g/L,其对美 洲斑潜蝇取食、产卵拒避持效期分别是4-8天和10天。使用浓度0.005 g/L揭阳霉素处理6天、8天和10天后接虫,幼虫的存活率分别是0、16.13%和28.07%。田间使用浓度0.005 g/L的揭 阳霉素和0.0045 g/L几的害极灭分别处理,6天后的校正虫口减退率分别为91.0%和90.9%,两者差异不显著,而使用浓度0.0067 g/L揭阳霉素处理,6天后校正虫口减退率为93.6%。     

Polyandry and Reproduction in the Serpentine Leaf Miner Liriomyza trifolii (Diptera: Agromyzidae)

We investigated the hypotheses that Liriomyza trifolii is a polyandrous species and that multiple mating is required to maximize female fecundity. In addition, the hypothesis that male copulating frequency reduces female fecundity was examined. Using sterile and fertile males, we found that L. trifolii females mate more than once during their life span, and that multiple mating is required to maximize female fecundity. A similar amount of sperm was transferred during five successive male copulations with an equal insemination probability. However, the probability of reproduction failure was strongly correlated with the male copulating sequence. Moreover, the failure rate of sperm transfer was found to be 6.8%. In light of these results, possible selection pressures driving for polyandry in L. trifolii are discussed.     

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.     

美洲斑潜蝇幼虫空间分布型和田间抽样技术

通过田间调查和计算,明确了美洲斑潜蝇幼虫呈聚集分布,且以负二项分布为主,理论抽样数当t＝1．00,D＝0．2时,n＝26．08／X＋16．12,如果防治指标定为10头／株,则最大抽样数为19株,序贯抽样的累积幼虫数量界限为．田间随机取样以平行线和Z字形为最佳。     

Variation of density-dependence with spatial scale in the leaf-mining fly Liriomyza commelinae (Diptera: Agromyzidae)

Abstract. 1. Mortality imposed on the leaf mining fly, Liriomyza commelinae (Frost) was studied at three spatial levels: (1) the leaf, (2) the stem, and (3) a metre quadrat of the food plant.
2. Real mortality (R.M.) (death as a percentage of eggs laid) during development for five sites over 3 years (forty-seven generations) was 96.3 + 1.9% -2.5% (±95% c.l.), due mainly to parasitoids and predatory ants, with a small component due to competition and other causes.
3. Real mortality due to the parasitoid complex averaged 41.2%. It was not related to population density in leaves or stems but was positively dependent on density in one metre quadrats.
4. Predation by Crematogaster brevispinosa Mayr was 21.2% and was independent of population density at all spatial levels.
5. Mortality due to other causes (e.g. competition and exposure to high temperature resulting from insolation) was positively dependent on the number of mines per leaf at three out of five sites studied and the number of mines per stem at two of the three sites studied; however, such mortality was independent of density per metre quadrat.     

Comparative demography of Liriomyza sativae Blanchard （Diptera： Agromyzidae） on cucumber at seven constant temperatures

Reproduction and population parameters of vegetable leafminer, Liriomyza sativae Blanchard were measured on cucumber （Cucumis sativus L.） at seven constant temperatures （10, 15, 20, 25, 30, 35 and 40℃）. No eggs were found at 10℃ and flies died after exposure to 40℃. The significantly highest intrinsic rate of natural increase （rm）, net reproductive rate （R0） and finite rate of increase （λ） ofL. sativae were obtained at 25℃ as 0.196, 52.452, and 1.216, respectively. The above-mentioned parameters decreased at 15℃ and 135℃ and this reduction at 35℃ was strong. Doubling time （DT） varied significantly with temperature. The shortest doubling time was obtained at 25℃. Mean generation time （T） decreased significantly with increasing temperature between 15℃ and 35℃. Percentage of immature ages in the stable age distribution was more than 95% at all temperatures. Female longevity was greater than male at all temperatures. Liriomyza sativae lived for a long time at 15℃, whereas at 35℃ had lower survival rates. The effect of temperature on reproduction, especially the intrinsic rate of increase of L. sativae would be useful for predicting its longterm population fluctuation over several generations.     

The feasibility of using the sterile insect technique against Liriomyza trifolii (Diptera: Agromyzidae) infesting greenhouse chrysanthemum

Liriomyza trifolii is a serious pest of chrysanthemum greenhouses and other crops around the world. The larvae feed within the leaves of the host plants and create serpentine mines. The Sterile Insect Technique (SIT) has been successfully used against some Dipteran species in the field, and it is especially efficient against isolated populations like those on islands. Therefore, SIT against pest populations in confined environments such as greenhouses should have high potential for success. The objectives of this study were to determine the optimum gamma‐irradiation dosage required for the sterilisation of L. trifolii adults, and to determine the quality (emergence percentage, flight ability, longevity, copulatory success, and sperm transfer) of the irradiated males compared with that of non‐irradiated (normal) males. We found that sterility (< 0.7 mine per female) was achieved with the dose of 170 ± 5% Gy. The copulatory success and sperm transfer during copulation of the sterile males were not significantly different from those of normal males. Moreover, the longevity, emergence percentage and flight ability of irradiated males were also not significantly different from normal males. The SIT experiments suggest that the release of sterile L. trifolii can significantly reduce the reproductive capacity of a wild leafminer population. Our studies indicate that sterilisation of L. trifolii flies is feasible and that sterile males are of high quality and competitive with normal males. Based on these data, research on the use of SIT against L. trifolii populations in greenhouses is ongoing.     

Role of plant volatiles in host plant location of the leafminer, Liriomyza sativae (Diptera: Agromyzidae)

Abstract The role of plant volatiles in host plant location of the leafminer Liriomyza sativae Blanchard was studied. Four types of antennal sensilla were identified on the funiculus by scanning electron microscopy: trichoid, basiconic, clavate and grooved sensilla. An olfactory pit, containing groups of sensilla, was present on the ventral side of the funiculus. No sexual difference was detected in sensilla diversity and distribution. In behavioural assays, both males and females were attracted by the odour of the bean Phaseolus vulgaris L. They had distinct EAG responses to the bean odour. No significant sexual difference was found in behaviour or EAG responses.
Electroantennograms were recorded from female L. sativae to 14 plant volatile compounds. The most distinct EAG responses were obtained for: (1) the general green leaf volatiles 1-hexanol (E)-2-hexen-1-ol, (E)-3-hexen-1-ol and its isomers, (Z)-3-hexen-1-ol, the acetate (E)-3-hexenylacetate and the aldehyde hexanal; and (2) limonene, a compound associated with tomato, which is a key host plant of this insect. Other volatile compounds associated with host plants, such as α-pinene, myrcene, β-caryophyllene, and eugenol did not elicit responses. The ability of this insect to locate a host plant appears to be augmented by the perception of a combination of host-specific and general green leaf volatiles. A modification of the EAG recording method of Dipteran species was provided.     

Effects of Melia azedarach, (Meliaceae) fruit extracts on the leafminer Liriomyza huidobrensis, (Diptera, Agromyzidae): Assessment in laboratory and field experiments

Control of the widely distributed pest Liriomyza huidobrensis, is complicated due to the protected habit of the leafmining larvae, and their resistance to insecticides. The effects of Melia azedarach, (Meliaceae) fruit extracts against adults and larvae of L. huidobrensis, were investigated. In the laboratory, leaves of Cucurbita, sp. infested with first and third instar larvae were treated with different extract solutions. Larval and pupal survival, as well as wing‐spread of adults, were ssessed. Female adult behaviour towards the extract was also analysed in terms of number of feeding punctures and number of offspring left on treated/untreated leaves. In the field, an infested Vicia faba, crop was sprayed four times at weekly intervals with plant extract, water, and a blank solution. The number of adult leafminers and parasitoids emerging from sampled leaves from each treatment were compared. The laboratory tests showed translaminar action of the extracts, which negatively affected leafminer pupal survival, while body size was not affected. The extracts also deterred feeding by adult females and may also have caused reduction in oviposition rates. All solutions and concentrations tested had similar effects. In the field, extract effects were consistent with those from laboratory trials, number of pupae and pupal survival being lower on treated plants. Percentage parasitism was not affected by plant extract treatment, suggesting a selective activity.     

The separation of Liriomyza huidobrensis (Diptera: Agromyzidae) from related indigenous and non-indigenous species encountered in the United Kingdom using cellulose acetate electrophoresis

The non-indigenous pest leaf miner Liriomyza huidobrensis and its close relatives L. bryoniae and L. strigata belong to a natural group. These species can be separated from one another by means of protein electrophoresis on a cellulose acetate membrane. The protocol for this diagnostic assay involves staining for two enzymes, glucose-6-phosphate dehydrogenase (G6PDH) and leucine-glycine peptidase (PEP). Other leaf miner species that might also be found under glass, Chromatomyia syngenesiae, C. horticola and the non-indigenous L. trifolii and L. sativae , are clearly distinguishable from the L. huidobrensis group of species by this assay. The effect of parasitism on L. huidobrensis by Dacnusa sibirica was investigated and shown unlikely to cause misidentification of the leaf miner host. Results obtained from practical use of the assay to identify unknown specimens from ports and nurseries are compared with those obtained from flies kept in laboratory culture and used during development of the assay. No new variation was found for G6PDH. New PEP variation was found for both L. huidobrensis and L. bryoniae but this did not affect the integrity of the assay. A biochemical key to these agromyzid leaf miners is presented.     

不同地理种群美洲斑潜蝇及近缘种的rDNA-ITS1序列分析和比较

【目的】通过对美洲斑潜蝇Liriomyza sativae Blanchard 不同地理种群及近缘种间的核糖体DNA第一内转录间隔区（rDNA-ITS1）进行比较,分析美洲斑潜蝇不同地理种群间的遗传分化情况,并为美洲斑潜蝇与近缘种间提供分子鉴别标记。【方法】用PCR产物直接测序法及克隆测序法对我国美洲斑潜蝇8个地理种群的rDNA-ITS1序列进行测序,并调用GenBank中3个近缘种的rDNA-ITS序列,运用软件MEGA3.1对美洲斑潜蝇不同地理种群及近缘种间的rDNA-ITS1序列进行分析。【结果】美洲斑潜蝇8个地理种群间的分化程度较低,只有8个变异位点,遗传距离都在0.02以下,但4个近缘种间的碱基差异显著,遗传距离为0.149～0.390,有126个变异位点,12个美洲斑潜蝇特异性识别位点。【结论】虽然基于rDNA-ITS1序列所显示的美洲斑潜蝇各地理种群之间的遗传分化很小,但是其分化趋势与地理分布基本相吻合;得到的12个特异性识别位点不仅可以作为美洲斑潜蝇与其近缘种间鉴别的分子标记,而且可为今后设计鉴别性PCR引物提供重要的参考依据。     

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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Molecular phylogeny of Chinese Rubia (Rubiaceae: Rubieae) inferred from nuclear and plastid DNA sequences

Rubia L. is the type genus of the coffee family Rubiaceae and the third largest genus in the tribe Rubieae, comprising ca. 80 species restricted to the Old World. China is an important diversity center for Rubia, where approximately half of its species occur. However, its internal phylogenetic relationships are still poorly understood. The objective of the present study is to contribute to the phylogenetic relationships within Rubia, using the nuclear internal transcribed spacer and six plastid markers and focusing on species from China. Twenty-seven species of Rubia were sampled to infer their phylogeny using Maximum parsimony, Maximum likelihood, and Bayesian analyses. The monophyly of Rubia is supported, provided that R. rezniczenkoana Litv. is excluded from Rubia and transferred to Galium as a new combination: G. rezniczenkoanum (Litv.) L. E Yang & Z. L. Nie. Within Rubia, two clades are clearly supported. They correspond to the traditional sect. Rubias.l. (A) and sect. Oligoneura Pojark. (B), and are morphologically mainly separable by their pinnate (A) versus palmate (B) leaf venation. Plesiomorphic features are the pinnate leaf veining in sect. Rubia s.l. and the occurrence of some species with opposite leaves and true stipules in sect. Oligoneura. In sect. Oligoneura one can assume an evolution from species with opposite leaves and true stipules (as in the R. siamensis Craib group) to those with whorls of two leaves and two leaf-like stipules (as in ser. Chinenses and the R. mandersii Collett & Hemsl. group) and finally with whorls of 6 or even more elements (as in ser. Cordifoliae). The correlation between clades recognized by DNA analyses and available differential morphological features is partly only loose, particularly in the group of R. cordifolia with 2×, 4×, and 6× cytotypes. This may be due to rapid evolutionary divergence and/or hybridization and allopolyploidy.     

三种寄主植物叶片提取物对美洲斑潜蝇雌成虫嗅觉记忆的影响

先提供气味刺激,再提供含不同寄主植物叶片提取物的蜜糖刺激,采用Y型嗅觉仪测定了3种寄主植物（丝瓜、番茄和菜豆）叶片表面非挥发性次生化合物刺激对美洲斑潜蝇Liriomyza sativae雌成虫嗅觉记忆的影响。结果表明：菜豆叶片沸水提取物能促进雌成虫对芳樟醇的嗅觉记忆,而丝瓜和番茄叶片沸水提取物则能提高雌成虫对正己烷的记忆能力。因此,不同寄主植物叶片提取物影响了美洲斑潜蝇雌成虫对不同气味的嗅觉记忆。     

南美斑潜蝇为害对黄瓜体内主要营养物质、 次生代谢物质及叶绿素含量的影响

南美斑潜蝇Liriomyza huidobrensis (Blanchard)是一种危害多种蔬菜和观赏植物的多食性害虫。本研究采用蒽酮比色法、 考马斯亮蓝法、 磷钼酸 磷钨酸比色法、 索氏回流法及丙酮法, 分别测定了南美斑潜蝇1~3龄幼虫不同为害程度（系统受害、 轻度受害和重度受害）对黄瓜叶片内可溶性糖、 可溶性蛋白质、 单宁、 黄酮及叶绿素含量的影响。结果表明： 南美斑潜蝇幼虫为害后, 黄瓜叶片中可溶性糖、 可溶性蛋白质和叶绿素的含量随着为害程度的增强而显著降低（P<0.01）, 最大降幅分别为62%, 35%和40%; 单宁和黄酮的含量随着为害程度的增强而显著上升（P<0.01）, 最高升幅分别为26%和53%; 并且具有系统性影响。结果提示, 南美斑潜蝇幼虫为害导致寄主植物营养物质含量下降、 光合作用降低, 而次生代谢物质含量上升, 从而对植食者产生诱导抗性。本研究结果为进一步揭示南美斑潜蝇与其寄主植物相互作用关系及其机理提供了有益的参考。     

温度对美洲斑潜蝇及南美斑潜蝇种群增长的影响

在恒温条件下观察了美洲斑潜蝇和南美斑潜蝇实验种群的发育、存活及繁殖情况,拟合了两种斑潜蝇发育速率与温度之间的关系模型,求出了各发育阶段的发育起点温度及有效积温,组建了不同温度下两种斑潜蝇实验种群的生殖力表,计算了主要的生命表参数.结果表明,温度对两种斑潜蝇的发育、存活、繁殖均有显著的影响,且两种斑潜蝇对温度的适应特性有着明显差异.美洲斑潜蝇适应的温度范围较广,相对较高的温度有利于种群的发育、生存及繁殖;而南美斑潜蝇适应的温度范围相对较窄,且适温范围明显偏低,高温不利于种群的增长;在各自的适温条件下,两种斑潜蝇都有很强的种群增殖能力.