Age-dependent response of female melon fly,Zeugodacus cucurbitae (Diptera: Tephritidae), to volatiles emitted from damaged host fruits

In many insect species including fruit flies (Diptera: Tephritidae), the behavioral responses to volatiles emitted by their host plants can be modulated by environmental conditions and by the physiological state of the insect. Here, we quantified (1) the effects of female age on the attraction of female melon fly, Zeugodacus cucurbitae (Diptera: Tephritidae) to volatiles emitted by intact and mechanically damaged tomato (Solanum lycopersicum), eggplant (S. melongena), zucchini (Cucurbita pepo), bitter melon (Momordica charantia), and cucumber (Cucumis sativus) fruit, and (2) the influence of time elapsed since fruit damage on the outcome. The investigations were conducted under semi-natural conditions in Hawai’i. Results from the first experiment revealed that, for freshly damaged tomato, eggplant, and zucchini, the level of female response was comparatively low and was not affected by female age. For bitter melon and cucumber, higher levels of response to freshly damaged fruit were documented, and the response levels gradually increased as female age increased from 1 to 4 weeks, reaching 70% for 4-week-old females exposed to cucumber odor. Results from our second experiment indicated that, on average, 56% of the females released responded in 20 min when cucumbers were freshly sliced, and the level of response was reduced 6-fold within a couple of hours. Females did not respond to tomato and cucumber odor when fruits were damaged >8 h before testing. Fruit volatiles involved in female Z. cucurbitae attraction seem to be released shortly upon mechanical damage and they are short-lived. The plasticity of the olfaction-driven behavior observed in Z. cucurbitae depending on female age and on the strength of the olfactory cues associated with preferred and less preferred hosts, adds another dimension to our understanding of the host-seeking behavior of this invasive species.     

Trapping tephritid fruit flies (Diptera: Tephritidae) in citrus groves of Fujian Province of China

Certain tephritid fruit flies, such as the oriental fruit fly, Bactrocera dorsalis, the Chinese citrus fly, B. minax and the Japanese orange fly, B. tsuneonis (Diptera: Tephritidae: Dacinae), are destructive citrus pests in China. A two-year trapping study was conducted in pomelo, Citrus maxima, groves in Fujian Province of China. The objectives of this study were to investigate the species, the abundance of tephritid fruit flies in the orchards, as well as the efficacy of the selected lure traps to these flies. Four lure traps or devices, i.e. methyl eugenol + Steiner trap (ST), cuelure + ST, ammonium acetate + putrescine + ST, and sticky spheres, were deployed from June to November 2017 and April to October 2018. Six economically significant Dacini pests were trapped during the period. These flies are B. dorsalis, the melon fruit fly, Zeugodacus cucurbitae, the pumpkin fruit fly, Z. tau, the Malaysian fruit fly, B. latifrons, and other two species - B. rubigina and Z. scutellatus. B. dorsalis was the most abundant, accounting for more than 50% of the capture, followed by Z. cucurbitae. The remaining four species accounted for less than 2% of the total capture. B. minax and B. tsuneonis, two destructive citrus-damaging tephritid fruit flies in China, were not found during the trapping period. Methyl eugenol trapped the highest number of fruit flies, followed by cuelure.     

Fruit flies reared from Terminalia catappa in Thailand

Bactrocera fruit flies (Diptera: Tephritidae) are pests of cultivated plants worldwide. Many Bactrocera flies are specific to commercial vegetable and fruit crops but some may develop in alternate hosts. One such alternate host is malabar or Indian almond, Terminalia catappa L. (Combretaceae) in Thailand. We studied a wild mature tree that was at least 20 years old and growing in a protected environment of Kasetsart University Kamphaeng Saen campus in 2008. We found that fruit flies and their associated parasitoids were recovered only from ripening (yellow colored) malabar almond fruits. Four species of fruit flies were recorded: Bactrocera dorsalis (Hendel), B. correcta (Bezzi), B. latifrons (Hendel) and B. cucurbitae (Coquillett). Of the four species of parasitoids recorded; the braconid Fopius arisanus (Sonan) (Hymenoptera: Braconidae) was most commonly recovered.     

Genetic structure and demographic history of the melon fly Zeugodacus cucurbitae (Coquillet) (Diptera: Tephritidae) in Thailand

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Oviposition acceptance and larval development of Chilo partellus stemborers in drought‐stressed wild and cultivated grasses of East Africa

Maternal host choices during oviposition by herbivorous insects determine the fitness of their offspring and may be influenced by environmental changes that can alter host‐plant quality. This is of particular relevance to ‘push‐pull’ cropping systems where host preferences are exploited to manage insect pest populations. We tested how drought stress in maize and companion plants that are used in these systems affect oviposition preference, larval feeding, and development of the spotted stemborer, Chilo partellus Swinhoe (Lepidoptera: Crambidae). Five host species were tested (all Poaceae): maize (Zea mays L.), Napier grass (Pennisetum purpureum Schumach), signal grass [Brachiaria brizantha (A. Rich) Stapf], Brachiaria cv. ‘Mulato’, and molasses grass [Melinis minutiflora (Beauv.)]. Under drought stress, maize experienced as much oviposition as control unstressed maize in choice and no‐choice experiments. Similarly, larval leaf damage was not significantly different in drought‐stressed vs. unstressed maize. In contrast, oviposition occurred less on drought‐stressed than on unstressed Napier and signal grass. Oviposition acceptance and leaf damage remained low in both drought‐stressed and unstressed molasses grass and Mulato. Larval survival and development remained high in drought‐stressed maize, but not in Napier, signal, and molasses grass and Mulato, where survival and development were low in both drought‐stressed and unstressed plants. Our results indicate that herbivore responses to drought‐stressed plants depend on the plant species and that drought stress can change host preference and acceptance rankings. In particular, trap‐crops such as Napier grass may not divert oviposition from the main maize crop under drought stress conditions.     

Raspberry ketone-supplemented diet has no effect on fitness parameters or lure responsiveness in male melon flies,Zeugodacus cucurbitae (Diptera: Tephritidae)

The male annihilation technique (MAT) and sterile insect technique (SIT) are often used to control pestiferous tephritid fruit flies (Diptera: Tephritidae). MAT involves the deployment of traps containing a male attractant and insecticide with the goal of drastically reducing male abundance and ultimately eliminating the entire population. SIT, which involves the mass production, sterilization, and release of the target species, may also be implemented to achieve final extirpation. Generally, simultaneous implementation of MAT and SIT is counterproductive, because the presence of large numbers of male-specific traps in the environment (MAT) would greatly reduce the number of sterile males available for copulating with wild females (SIT). However, studies on the Queensland fruit fly, Bactrocera tryoni (Froggatt), indicate that concurrent use of MAT and SIT may be feasible. Sexually mature males of B. tryoni are attracted to the raspberry ketone and its synthetic analogue cue-lure. Males of B. tryoni fed raspberry-ketone-supplemented diet when newly emerged showed lower attraction to cue-lure baited traps than control males. In addition, newly emerged males provided this diet displayed accelerated sexual maturation, which would allow the early release of sterile males and reduce pre-release holding costs. Here, we examined whether the addition of raspberry ketone to the adult diet of male melon flies, Zeugodacus cucurbitae (Coquillett), produced effects similar to those observed for B. tryoni. Despite using similar methods, no significant effect of raspberry ketone-supplemented diet on time to sexual maturity, survival, mating competitiveness, or attraction to cue-lure baited traps in mass-reared Z. cucurbitae males.     

Capture of Bactrocera fruit flies (Diptera: Tephritidae) in traps baited with liquid versus solid formulations of male lures

Bactrocera dorsalis (Hendel) and B. cucurbitae (Coquillett) (Diptera: Tephritidae) are important agricultural pests of the Pacific region. Detection and control of these species rely largely on traps baited with male-specific attractants (parapheromones), namely methyl eugenol for B. dorsalis and cue lure for B. cucurbitae. Presently, these lures (plus naled, an insecticide) are applied in liquid form, although this procedure is time-consuming, and naled as well as methyl eugenol may pose human health risks. Recently, a solid formulation (termed a wafer) has been developed that contains both male lures (plus DDVP, an insecticide), and here we present data from field tests in California and Hawaii that compare the effectiveness of liquid versus solid formulations of the lures in capturing marked, released males of these two Bactrocera species. For both species and in both California and Hawaii, traps baited with the solid formulation of the male lure captured similar or significantly more released flies than the liquid formulation for both fresh and aged baits. Traps in Hawaii also captured wild (unmarked) males of both B. dorsalis and B. cucurbitae, and the results obtained for wild flies were similar to those recorded for released flies for both species. Collectively, the results presented suggest that the solid dispenser of the male lures constitutes a reliable substitute for the liquid formulation in detecting incipient Bactrocera outbreaks.     

Capture of melon flies, Zeugodacus cucurbitae (Diptera: Tephritidae), in a food-baited Multilure trap: Influence of distance,diet, and sex

Many countries operate trapping programs to detect invasions of pestiferous fruit fly species (Diptera: Tephritidae). Surveillance relies heavily on traps baited with male lures, which, while highly attractive, have limited effectiveness, because (i) they are sex-specific and (ii) males of some species do not respond to the lures currently in use. For these reasons, detection programs also include food-baited traps that are neither sex- nor species-specific. Compared to male lure-baited traps, however, few studies have measured the attractiveness of food-based traps. The present study describes a mark-release-recapture study conducted in a fruit orchard in Hawaii that measured the attractiveness of a liquid protein hydrolysate-based (torula yeast/borax slurry) trap to adults of the melon fly Zeugodacus cucurbitae (Coquillett). Multiple release points were used at varying distances from a single, central trap to generate estimates of distance-dependent capture probabilities. The potential influences of sex and pre-release diet on capture probability were also examined. Flies were released at 14 d of age and were maintained on one of four dietary regimes that offered a protein hydrolysate-rich diet for varying intervals (i.e., 0, 3, 7, or 14?d, respectively). Recapture rates were similar between the sexes and over both sexes and all diets averaged 3.6%, 3.2%, and 0.6% for release distances of 10, 25, and 50?m, respectively. Pre-release diet had a significant effect on recapture probability for releases at 10 and 25?m: flies fed sugar only or protein hydrolysate-rich diet for only 3?d were captured more frequently than flies that had longer access to yeast extract prior to release.     

Host plant selection behaviour of Chilo partellus and its implication for effectiveness of a trap crop

Female lepidopterans can display a hierarchy of preference among potential host species, a trait thought to arise from the balance between attractants and deterrents to which the insects respond. Host plant ranking by moths and larvae of Chilo partellus Swinhoe (Lepidoptera: Crambidae), an important pest of cereals in Africa, was investigated, and whether eggs deposited on specific host plants yield larvae of particular host preferences. Trap plants are used in management of this pest. However, any ‘disagreement’ in host ranking between moths and larvae could potentially reduce effectiveness of trap crops as larvae emigrate to the main crop from the parent’s preferred trap plant. We also investigated whether host plant preference is influenced by the diet upon which larvae fed as part of an integrated assessment of the relationship between host plant selection and learning in C. partellus. Five host plants (all Poaceae) were used: maize (Zea mays L.), sorghum (Sorghum bicolor Moench), Napier grass (Pennisetum purpureum Schumach), and two varieties of signal grass [Brachiaria brizantha (A. Rich.) Stapf], viz., local (henceforth signal grass) and improved (‘Mulato’). In multiple choice tests, C. partellus female moths preferentially oviposited on Napier grass, followed by sorghum, maize, and signal grass, and least preferred ‘Mulato’. Larvae however equally orientated and settled on leaf cuts of maize, sorghum, signal grass, and Napier grass, but least preferred ‘Mulato’. Moreover, eggs from specific host plants did not yield larvae of particular host preferences. Furthermore, oviposition preference was not altered by the larval food. These results imply only a slight ‘disagreement’ in host ranking behaviour between moths and larvae, which is beneficial for trap cropping as larvae would not ‘reject’ the trap plant and appreciably disperse to the neighboring plants. Moreover, absence of larval learning behaviour indicates that regardless of the larval food C. partellus moths would still be attracted to the selected trap plant.     

Synergistic and additive interactions among components of food-based baits underlie female fruit fly attraction

Attraction of tephritid fruit flies to some food sources can be enhanced by the presence of ammonia derivatives, compounds that are perceived as volatile cues for protein-rich food sources. Using a comparative approach, we (1) evaluated the behavioral responses of females of three invasive fruit fly species, Bactrocera (Zeugodacus) cucurbitae (Coquillett), Bactrocera dorsalis (Hendel), and Ceratitis capitata (Wiedemann) (all Diptera: Tephritidae) to trub (a protein-rich waste brewer’s yeast product generated during the production of beer), Concord grape juice (a protein-deficient material), and ammonium acetate, and (2) identified synergistic and additive interactions between low- and high-attractiveness materials and ammonium acetate. We established the attractiveness of fresh trub, grape juice, and ammonium acetate when tested singly to females of all three fly species. Although ammonium acetate did not enhance significantly the response of females of any species to fresh trub, the most attractive material, ammonium acetate, did significantly enhance females’ level of response to aged trub (a comparatively less attractive material) and to grape juice. Our research found a synergistic interaction between diluted grape juice and ammonium acetate for B. cucurbitae, as well as between aged trub and ammonium acetate for B. dorsalis. For C. capitata, additive effects among food attractants and ammonium acetate were identified. Our findings increase our understanding of fruit fly female olfactory-driven behavior in response to food-based materials and the extent to which ammonium acetate modulates female response to protein-rich and protein-deficient materials.     

地中海实蝇及其近缘种基因芯片检测研究

本研究选择线粒体DNA (mtDNA) 细胞色素氧化酶Ⅰ基因（COⅠ）为分子标记基因,以双翅目实蝇科昆虫DNA序列为目标,建立了我国进境植物检疫害虫地中海实蝇Ceratitis capitata、芒果小条实蝇C. cosyra和纳塔尔小条实蝇C. rosa等生物芯片检测方法。地中海实蝇及其近缘种检测芯片由检测探针（实蝇科通用探针1条,小条实蝇属通用探针1条,地中海实蝇、芒果小条实蝇和纳塔尔小条实蝇近缘种探针2条和种特异探针4条）、质控探针（定位点探针、阳性质控、阴性质控和空白对照探针各1条）组成。芯片检测结果表明,检测探针特异性强,能实现上述3种实蝇的种类快速区分和准确鉴定; 检测方法稳定性好,地中海实蝇不同虫态（卵、幼虫、蛹和成虫）和不同地理种群检测结果完全一致。地中海实蝇生物芯片检测技术将为我国进口果蔬中检疫性实蝇快速筛查和种类鉴定提供检测方法,同时,还可应用到其他属的实蝇以及相关害虫的检疫中,为有害生物的快速鉴定提供了新方法。     

Population structure of the melon fly, Bactrocera cucurbitae, in Reunion Island

The melon fly, Bactrocera cucurbitae (Coquillett) (Diptera: Tephritidae) is an agricultural pest of major significance worldwide that primarily attacks cucurbit crops. In Reunion Island, it represents the main tephritid pest on cucurbits. In this paper, we provide a genetic characterization of populations of B. cucurbitae from Reunion Island and investigate their geographical origin using ten microsatellite loci at two mitochondrial gene fragments. Microsatellites reveal the occurrence of three different genetic clusters of B. cucurbitae in Reunion Island, all clearly distinguishable from their African and Asian relatives. These three clusters are sympatric and show no signs of recent bottlenecks. Levels of gene flow among clusters are relatively high, yet gene flow also occurs with populations from the African continent and, to a lesser extent, from Asia. The B. cucurbitae clusters show distinct distributions across eastern and western locations in Reunion Island (but not at different altitudes or between wild and cultivated host plants or between sampling periods), and their abundance is also correlated with the average amount of rainfall. Microsatellite and sequence analyses suggest Africa as the most probable source area for populations of B. cucurbitae in Reunion Island.     

Flight and oviposition behaviour of the African stem borer,Busseola fusca,on various host plant species

The African stem borer, Busseola fusca (Fuller) (Lepidoptera: Noctuidae), is an important pest of maize and sorghum in sub‐Saharan Africa. As in many other lepidopteran insects, the ability of B. fusca to recognize and colonize a variety of plants is based on the interaction between its sensory systems and the physical and chemical characteristics of its immediate environment. In this study, we tried to identify the behavioural steps of B. fusca leading to host selection and oviposition. Three Poaceae species commonly cultivated in Kenya for human consumption and animal forage were used in this study: the two most preferred hosts, maize (Zea mays L.) and sorghum [Sorghum bicolor (L.) Moench], and one non‐preferred host, Napier grass (Pennisetum purpureum Schumach). Wind tunnel observations revealed that volatiles produced by the different plant species did not appear to strongly influence the general orientation of B. fusca towards the plant, as similar behavioural steps were exhibited by the female moth regardless of the plant species involved. This indicated that the females were not able to recognize their preferred hosts from a distance. After landing, the female typically swept her ovipositor on the plant surface, simultaneously touching it with the tips of her antennae, and then oviposited. This behaviour was more frequently observed on maize and sorghum than on Napier grass, and indicated that both antennal and ovipositor receptors are used by the female moths to evaluate the plant surface before deciding to oviposit. As a result, the females laid more eggs on the two crops than on Napier grass. We conclude therefore that females recognized their preferred hosts only after landing. Tactile and contact‐chemoreception stimuli from the plants seemed to play a major role in oviposition decisions of B. fusca.     

γ‐Octalactone,an effective oviposition stimulant of Bactrocera tryoni

Insects commonly rely on olfactory, gustatory and visual cues when deciding where to lay eggs. The olfactory cues that stimulate oviposition in the Queensland fruit fly, Bactrocera tryoni (Froggatt) (Diptera: Tephritidae), are not well understood. Here, we show that two known oviposition stimulants of the Oriental fruit fly, Bactrocera dorsalis (Hendel) (Diptera: Tephritidae)—γ‐octalactone and benzothiazole—strongly elicit aggregation and oviposition in B. tryoni. Two other known oviposition stimulants of B. dorsalis—ethyl tiglate and 1‐octen‐3‐ol—elicit aggregation but not oviposition. Highlighting species overlap, but also differences, in oviposition stimulants, these findings have practical application for mass‐rearing in which vast numbers of flies are reared for sterile insect technique programs and may also have practical application in the development of pest management and monitoring tools.     

云南主要有害实蝇种类及区划

采用诱剂诱捕辅以受害瓜果调查的方法,系统梳理了云南有害实蝇类群,确定主要有害种类25种。基于云南自然地理特征,采用聚类分析法深入探讨了云南实蝇区系,提出云南实蝇地理区划可分为德宏、版纳盆谷区,南部边缘中低山宽谷区,滇西南山原峡谷区,中东部高原、岩溶山原区以及北部高原山地区等5大区域。研究发现,受热量条件制约,云南主要有害实蝇类群呈南北向梯度递减;受纵向岭谷区"通道-阻隔"综合作用影响,云南西南部河谷地区是实蝇危害较为严重的区域。首次系统揭示了云南主要有害实蝇的地理分布规律。     

Capture of Bactrocera fruit flies in traps baited with liquid versus solid formulations of male lures in Malaysia

Approximately 70 species of Bactrocera fruit flies (Diptera: Tephritidae) are polyphagous economic pests that attack many important agricultural crops. Several of these Bactrocera species are also highly invasive, and many countries operate continuous, large-scale trapping programs to detect incipient infestations. Detection programs rely heavily on traps baited with male lures, with males of some species responding to raspberry ketone (RK; or its synthetic analogue cue-lure [CL]) and males of other species responding to methyl eugenol (ME). These lures (plus naled, an insecticide) are currently applied as liquids, although this procedure is time-consuming and may expose workers to health risks. Recent field tests, conducted largely in Hawaii, have shown that traps baited with a solid formulation (termed a wafer) that contains both RK and ME (plus dichlorvos, an insecticide) capture as many or more B. dorsalis (Hendel) and B. cucurbitae (Coquillett) males as traps baited with the standard liquid lures. While these results are promising, a more complete evaluation of the solid formulation requires testing in a region with a diverse assemblage of Bactrocera species, since interspecific variation in male response to lures has been reported. The objective of the present investigation was to assess the relative effectiveness of liquid versus solid formulations of male lures in Malaysia, a country known to harbor a large assemblage of Bactrocera species. Based on a 12-week sampling period, we found that, contrary to the Hawaiian results, traps baited with the wafer captured significantly fewer males than traps baited with liquid lures for all five ME-responding taxa analyzed and for one of the three RK/CL-responding species analyzed. Possible explanations for the discrepancy between these and earlier findings are offered.     

Selection of non‐target tephritids for risk evaluation in classical biocontrol programmes against the olive fruit fly

The olive fruit fly, Bactrocera oleae (Rossi) (Diptera: Tephritidae), is the key pest of olives (Olea europaea L.). Classical biological control against this insect was previously attempted in Spain with Psyttalia concolor (Szépligeti) (Hymenoptera: Braconidae), with very limited success. Other parasitoids are now available for new classical biological control programmes. Before release of exotic parasitoids, an assessment of their potential impact on non‐target species is required. Surveys were conducted in different olive groves in south‐eastern Madrid to study wild tephritids associated with Asteraceae plants. We recorded plant species and their abundance and collected flower heads to identify and quantify tephritid species. Fruits from Rosa canina L. were also collected. After a multicriteria analysis (MCA), we propose Urophora hispanica Strobl, U. stylata (Fabricius) and Carpomya schineri (Loew) as candidates for further risk assessment experiments. Additional information on new associations between tephritid flies and Asteraceae plants and on autochthonous Hymenoptera parasitizing tephritids is provided.     

Evaluation of Napier grass (Pennisetum purpureum) varieties for use as trap plants for the management of African stemborer (Busseola fusca) in a push–pull strategy

We evaluated eight Napier grass [Pennisetum purpureum Schumach (Poaceae)] varieties, used in various parts of eastern Africa as fodder, for their potential role as trap plants in the management of the African stemborer, Busseola fusca Füller (Lepidoptera: Noctuidae) through a push–pull strategy. Oviposition preference, larval orientation, settling, arrest and dispersal, feeding, mortality and survival, and development were determined for each of these varieties under laboratory and screen house conditions. Two‐choice tests showed that only two of the varieties tested (cv. Bana and cv. Uganda Hairless) were preferentially chosen by gravid female moths for oviposition over a susceptible maize variety, cv. Western Hybrid 502. Larval preference was, however, highly variable. Larval feeding by first instars on the maize leaves was more intense and significantly more than on leaves of all the Napier grass varieties evaluated. Food consumed and amounts assimilated by the third instars over a 24‐h period were not different among larvae fed on stems of maize and those fed on stems of the various Napier grass varieties. Larval survival was significantly lower on all the Napier grass varieties (below 3%) than on maize (about 44%). Similarly, larval development was about 2–3 weeks longer on majority of the Napier grass varieties. It was concluded that cv. Bana had potential for use as a trap plant in the management of B. fusca because it was more preferred by the moths for oviposition, equally preferred as maize by the larvae for orientation, settling, and arrest, and allowed minimal survival of the larvae. It can thus be used with such ‘push’ plants as Desmodium spp. (Fabaceae) in a ‘push–pull’ strategy, but the effectiveness of such a strategy would strictly depend on proper establishment and management of these companion plants.     

Lekking Behaviour and Male-Male Rivalry in the Melon Fly <Emphasis Type="Italic">Bactrocera cucurbitae</Emphasis> (Coquillett) (Diptera: Tephritidae)

Most studies on the sexual behaviour of tephritid fruit flies are conducted in laboratory or field cage settings. Here, we present field data on the mating behaviour of the melon fly, Bactrocera cucurbitae (Coquillett). A number of Tephritidae are lekking species, forming aggregations in which males fight to defend a small territory where they court females and mate. Our results confirm that males of the melon flies congregated and engaged in wing vibration in the late afternoon on non-host plants, such as Ficus carica, Cydonia oblonga and Prunus persica which bordered cucurbit fields. The nature and outcome of male-male aggressive encounters are described, and a high degree of female mate selectivity is documented.     

实蝇共生菌研究进展

实蝇类昆虫(双翅目Diptera:实蝇科Tephritidae)是重要的农业害虫,为害农作物茎秆和水果,严重影响农业生产和果蔬贸易,被许多国家列为重要的检疫性有害生物。实蝇共生菌对宿主实蝇的取食、生殖、发育及环境适应能力具有重要作用。目前,已有21个属的共生细菌(Enterobacter、Klebsiella、Citrobacter、Pseudomonas、Providencia、Erwinia、Acetobacter、Serratia、Proteus、Hafnia、Cedecea、Arthrobacter、Lactobacillus、Micrococcus、Streptococcus、Staphylococcus、Vibrio、Hafnia、Deinococcacea、Bacillus、Wolba-chia)以及1个属的共生真菌(Candida)被鉴定。其中,肺炎杆菌Klebsiella pneumoniae、产酸克雷伯氏菌Klebsiella oxytoca、成团泛菌Pantoea agglomerans、费氏柠檬酸杆菌Citrobter freundii、阴沟肠杆菌Enterobacter cloaca...