The tympanal hearing organ of the parasitoid fly Ormia ochracea (Diptera,Tachinidae, Ormiini)

Tympanate hearing has evolved in at least 6 different orders of insects, but had not been reported until recently in the Diptera. This study presents a newly discovered tympanal hearing organ, in the parasitoid tachinid fly, Ormia ochracea. The hearing organ is described in terms of external and internal morphology, cellular organization of the sensory organ and preliminary neuroanatomy of the primary auditory afferents. The ear is located on the frontal face of the prothorax, directly behind the head capsule. Conspicuously visible are a pair of thin cuticular membranes specialized for audition, the prosternal tympanal membranes. Directly attached to these membranes, within the enlarged prosternal chamber, are a pair of auditory sensory organs, the bulbae acusticae. These sensory organs are unique among all auditory organs known so far because both are contained within an unpartitioned acoustic chamber. The prosternal chamber is connected to the outside by a pair of tracheae. The cellular anatomy of the fly's scolopophorous organ was investigated by light and electron microscopy. The bulba acustica is a typical chordotonal organ and it contains approximately 70 receptor cells. It is similar to other insect sensory organs associated with tympanal ears.The similarity of the cellular organization and tympanal morphology of the ormiine ear to the ears of other tympanate insects suggests that there are potent constraints in the design features of tympanal hearing organs, which must function to detect high frequency auditory signals over long distances. Each sensory organ is innervated by a branch of the frontal nerve of the fused thoracic ganglia. The primary auditory afferents project to each of the pro-, meso-, and metathoracic neuropils. The fly's hearing organ is sexually dimorphic, whereby the tympanal membranes are larger in females and the spiracles larger in males. The dimorphism presumably reflects differences in the acoustic behavior in the two sexes.     

The tympanal hearing organ of a fly: phylogenetic analysis of its morphological origins

A key adaptation for any parasitoid insect is the sensory modality that it uses to locate its host insect. All members of the speciose family Tachinidae (Diptera) are parasitoids, but only flies of the tribe Orminini use acoustic cues to find their hosts. Ormiine flies are parasitoids of various genera of crickets and katydids. Gravid females of one ormiine species, Ormia ochracea, hear the reproductive calling song of male field crickets and home in on those calls to locate their hosts. While many flies possess various kinds of ears to detect airborne sounds, only ormiine flies have been reported to possess true tympanal hearing organs. Such organs are wellknown to occur in their cricket and katydid hosts. The ormiine ear is an evolutionary innovation within Diptera. Our objective was to trace the phylogenetic origins of the tympanal hearing organ among higher flies. Since the ormiine hearing organ is a complex organ within the prothorax, we examined possible precursor structures in the prothoraces of selected Diptera. We have uncovered a suite of characters that define the ormiine ear. These characters in the prothorax include a pair of prosternal tympanal membranes, a pair of chordotonal sensory organs, and modifications of the tracheal system. We have been able to identify and trace the presumptive homologs of these ormiine characters through selected species of related Diptera, using the method of outgroup comparison.Dedicated to the memory of Dr. Edmund A. ArbasThis work was supported by grants from NIH (5RO1 DC 00103), NIMH (IKOS MH01148-01), NSF (240-1879A) and Hatch (NYC-191403) to R.R.H. and the Swiss Science Foundation and the Janggen-Pöhn Foundation to D.R.     

Comparison of auditory sense organs in parasitoid Tachinidae (Diptera) hosted by Tettigoniidae (Orthoptera) and homologous structures in a non-hearing Phoridae (Diptera)

The dipteran parasitoids Therobia leonidei and Homotrixa alleni (Tachinidae) use acoustic cues to locate their calling tettigoniid (Ensifera, Orthoptera) hosts. The sexually dimorphic tympanal organs of both fly species are located at the prosternum. For comparison a homologous chordotonal organ in the non-hearing fly Phormia regina, Meigen (Phoridae) is also described. The scolopidial sense organs of the ears have approximately 180 sensory cells in Th. leonidei and 250 cells in H. alleni. Interspecific analysis indicates that the cell number and arrangement might be genus specific in Tachinidae. The mononematic scolopidia, each with one sensory cell, are of different sizes and insert at the tympanal membrane. Large scolopidial units (diameter of sensory cells up to 50 μm) extend longitudinally from the centre of the sensory organ towards the ligament, whereas small units (sensory cell diameter up to 10 μm) are arranged sequentially within the sensory organ. This arrangement is discussed to be a possible basis for frequency discrimination. The ultrastructure of the scolopidia is similar in the hearing and non-hearing flies. In both groups, the majority of scolopales has a diameter from 2 to 2.9 μm, although hearing species have additionally wider scolopales. The homologous chordotonal organ of Ph. regina consists of approximately 55 sensory cells of uniform direction. The data are discussed in comparison to the ears of other Diptera.     

中国瑟寄蝇属种类纪要（双翅目：寄蝇科）

研究了我国蜗寄蝇族瑟寄蝇属Cyrtophleba(双翅目:寄蝇科)2种,其中茹瑟寄蝇Cyrtophleba ruricola 分布于辽宁,山西,宁夏,新疆;变瑟寄蝇Cyrtophleba vernalis为中国新记录种,分布于辽宁。给出了上述2种的鉴别特征及变瑟寄蝇头和雄性尾器的特征图。研究标本分别保存于沈阳师范大学昆虫标本室(SYNU),中国科学院动物研究所(IZCAS)和国家林业局森林病虫害防治总站(GSFPM)。     

中国贵州长腹寄蝇属三新种记述（双翅目：寄蝇科）

记述采自贵州地区的长腹寄蝇属Dolichocoxys Townsend 3个新种:黄基长腹寄蝇D. flavibasis sp. nov.、黑腹长腹寄蝇D. obscurus sp. nov.和短柄长腹寄蝇D. brevis sp. nov.。文中附所有新种的详细描述、鉴别特征图、近缘关系的讨论及中国长腹寄蝇属分种检索表。新种的模式标本保存于贵州省安顺市疾病预防控制中心医学昆虫研究室。     

In Vitro Rearing of the Parasitoid Exorista larvarum (L.) (Diptera: Tachinidae) on Meat Homogenate-Based Diets

The tachinid Exorista larvarum (L.), a polyphagous gregarious larval endoparasitoid of Lepidoptera, was reared from egg to fecund adult on media containing commercial meat homogenates for babies as the main ingredient. Four media, each containing a diverse homogenate supplemented with extract of Galleria mellonella L. pupae, were tested first. Despite the difference in nutrient content, the kind of homogenate did not significantly affect the adult yields (30.2 to 40.7%) or puparial weights. Two other diets free of host materials (I and II) were then tested. Both were based on Gerber veal homogenate combined with different amounts of yeast extract and chicken egg yolk and were supplemented with wheat germ (I) or saccharose (II). Adult yields (28.7 to 32.7%) and puparial weights did not differ significantly between the two diets. Fly longevity and fecundity of the females obtained on diet I were comparable to those of the females emerged from puparia formed in G. mellonella larvae. Male and female puparial weights were, however, higher and development times longer on the diet than in the host.     

Reproductive biology of Fopius ceratitivorus (Hymenoptera: Braconidae), an egg–larval parasitoid of the Mediterranean fruit fly, Ceratitis capitata (Diptera: Tephritidae)

The reproductive biology of Fopius ceratitivorus Wharton, a recently discovered African parasitoid, was studied in quarantine in Hawaii to facilitate its mass production for biological control of the Mediterranean fruit fly, Ceratitis capitata. Mean longevity of host-deprived and ovipositing females was 17.3 ± 0.9 d and 16.2 ± 0.5 d, respectively. Ovarian maturation peaked at 61.6 mature eggs per female on the fifth day after eclosion and declined thereafter. Mean number of offspring produced per day by mated females was 5.1 ± 0.4, and realized fecundity expressed as total eggs deposited during the female’s life time was 107.8 ± 12.8. Females were more attracted, to and reproduced significantly more, in fruit substrates containing odors of adult flies and eggs rather than fruit substrates artificially inoculated with fly eggs. Our findings suggest that F. ceratitivorus is a promising new parasitoid for biological control of C. capitata in Hawaii.     

Response of a parasitoid fly, Gymnosoma rotundatum (Linnaeus) (Diptera: Tachinidae) to the aggregation pheromone of Plautia stali Scott (Hemiptera: Pentatomidae) and its parasitism of hosts under field conditions

We studied the attraction of a tachinid fly, Gymnosoma rotundatum (Linnaeus) to the male-produced aggregation pheromone of the brown-winged green bug, Plautia stali Scott, its parasitism on the bug, and its seasonal occurrence in the field. The tachinid fly was continuously attracted to the aggregation pheromone from spring to autumn and utilized the bugs as hosts. Our field experiment to clarify the effect of the pheromone on parasitism demonstrated that parasitism occurred only in female bugs baited with synthetic aggregation pheromone and did not occur in females without the pheromone. The parasitoid flies therefore appeared to use the bug’s pheromone as a host-finding kairomone. The pheromone attracted not only female flies but also males. Male flies may increase their chance of encountering pheromone-attracted females by waiting near pheromone sources. The tachinid develops multiple generations in active hosts from spring to autumn and overwinters in dormant hosts. Thus, G. rotundatum seems to be highly adapted to using P. stali as its host, and it is a potentially important biological control agent of P. stali populations in the field.     

Sexual dimorphism in the visual system of flies: The divided brain of male Bibionidae (Diptera)

Summary The mapping of the compound eyes onto the visual neuropils and the cell types in the lamina and the lobula complex of Bibionidae (Diptera) were studied by means of extracellular cobalt injections and Golgi impregnations. Dorsal and ventral eyes in males map into separate dorsal-and ventral neuropils up to the level of the lobula complex. The dorsal-eye lamina is unilayered, while the ventral-eye lamina in males and the lamina in females are multilayered: layers A and C are invaded by en-passant terminals of long visual fibres, layer B by the terminals of short visual fibres. Long visual fibres have a short and a long terminal in the ventral medulla with terminal specialisations in three distinct layers. Only one type of receptor ending exists in the dorsal medulla, the terminal branches of which are restricted to one layer only. Arrays of contralateral neurones are found in the medial part of the dorsal lobula, which receives input from the zone of binocular vision of the ipsilateral dorsal eye, and in the posterior dorsal lobula and lobula plate. The dorsal lobula plate contains large tangential neurones, the dendritic arborisations of which are revealed by cobalt injection into the thoracic ganglia. The divided brain of male bibionids offers the opportunity to investigate separately the nervous systems involved in sex-specific visually guided flight behaviour and in general visually guided flight control.     

温度对蝇蛹俑小蜂寄生瓜实蝇蛹功能反应的影响

【目的】蝇蛹俑小蜂是很多双翅目害虫的蛹寄生蜂。了解温度对寄生蜂寄生效能的影响是利用寄生蜂控制害虫的重要前提。【方法】实验室条件下设置温度为21、24、27、30、33℃,测定温度对蝇蛹俑小蜂雌蜂寄生瓜实蝇蛹功能反应的影响。蝇蛹俑小蜂对瓜实蝇蛹的功能反应用Michaelis-Menten-Ⅱ型功能反应模型N_a=AN/(F+N)进行拟合。【结果】不同温度下蝇蛹俑小蜂的寄生潜能(A)和寄主半饱和密度(F)不同。当温度在27℃时,其寄生潜能最大;而寄主半饱和密度随着温度升高而降低。【结论】瓜实蝇在瓜地盛发期的栖境温度适宜蝇蛹俑小蜂的寄生。     

Distribution, host specificity, and overwintering of Celatoria bosqi Blanchard (Diptera: Tachinidae), a South American parasitoid of Diabrotica spp. (Coleoptera: Chrysomelidae: Galerucinae)

The genus Diabrotica (Coleoptera: Chrysomelidae) includes a great number of pest species, including some of the most important crops pests of the Americas. However, only five parasitoid species have been recorded for it. The parasitoid Celatoria bosqi Blanchard was the first parasitoid described from Diabrotica spp. in South America, where substantial parasitism has been observed. C. bosqi has been collected almost throughout the South American distribution of its main host, Diabrotica speciosa (Germar), in an area that includes temperate and tropical lowlands, and semiarid to humid highlands. Three Diabrotica species were found to host the parasitoid, D. speciosa (Germar), Hystiopsis sp., and Diabrotica viridula (F.), with a total parasitism of 2.60, 5.55, and <0.02%, respectively. Laboratory experiments with field beetles and puparia, reared in the laboratory, indicate that C. bosqi overwinters obligatorily in overwintering adult host beetles, remaining quiescent in its live host below developmental temperatures. Based on the known climatic range of C. bosqi, and its requirement of adult overwintering hosts, a potential distribution in North America is projected.     

(Z)-3-nonenol, (Z,Z)-3,6-nonadienol and (S,S)-(-)-epianastrephin: male produced pheromones of the Mexican fruit fly

(Z)-3-nonenol, (Z,Z)-3,6-nonadienol and (S,S)-(-)-epianastrephin proved active as male-produced pheromones which elicited behavioral responses from virgin female Mexican fruit flies (Anastrepha ludens) (Diptera: Tephritidae) in laboratory bioassays. All three chemicals elicited attraction and/or locomotor arrest when tested individually. When tested together, (Z)-3-nonenol inhibited the behavioral effect of (Z,Z)-3,6-nonadienol but either of the alcohols synergized the effect of (S,S)-(-)-epianastrephin. Quantities of the pheromones per male abdomen were: (Z)-3-nonenol-100 ng; (Z,Z)-3,6-nonadienol-40 ng; and total epianastrephin (R,R and S,S enantiomers)-700 ng. Relative to the quantities in abdominal extract, males released these chemicals during sexual display in a blend containing a higher proportion of the alcohols.

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Résumé Le (Z)-3-nonènol, le (Z,Z)-3,6-nonadiènol et le (S,S)-(-)-épianastréphine produits par les mâles de la mouche mexicaine du fruit (Anastrepha ludens) (Diptera: Tephritidae) ont attirés les femelles lors de tests conduits en laboratoire. Le pouvoir attractif du (Z)-3-nonènol est faible comparé à celui des deux autres phéromones. Evalués ensemble, le (Z)-3-nonènol inhibe l'attractivité du (Z,Z)-3,6-nonadiènol mais chacun des deux alcools synergise l'effet attractif de la (S,S)-(-)-épianastréphine. Les quantités recouvrées par mâle dans un extrait abdominal étaient respectivement de 100, 40 et 700 ng pour le (Z)-3-nonènol, le (Z,Z)-3,6-nonadiènol, et l'épianastréphine totale (R,R et S,S) énantiomères. Comparées à ces valeurs, le mélange élange émis par les mâles lors de leur cour sexuelle est plus riche in alcools. Le rapport (Z)-3-nonènol (Z,Z)-3,6-nonadiènol est cependant identique dans l'extrait abdominal et dans le mélange émis par les mâles.

     

Sensory cell degeneration in the ontogeny of the chemosensitive sensilla in the labial palp-pit organ of the butterfly,Pieris rapae L. (Insecta,Lepidoptera)

Summary The ontogeny of the chemoreceptive sensilla in the labial palp-pit organ was studied in Pieris rapae by examining twelve successive stages between pupation and emergence of the imago, which takes a period of 160 h under the experimental conditions. Mitoses occur until 20 h after pupation. They lead to anlagen of sensilla, 91% of which are comprised of three sensory cells. However, two sensory cells degenerate in each sensillum during a period of 28 h. The same process occurs in anlagen with four sensory cells resulting in bicellular sensilla. Axons grow out only after the number of sensory cells has been reduced. Further consecutive steps in sensory cell differentiation are: (a) outgrowth of dendritic outer segment and dendrite sheath; (b) outgrowth of trichogen process and change in structure of elongating dendrite sheath; (c) deposition of cuticle and pore tubules in the pegs; (d) retraction of trichogen process; (e) increase in diameter of dendritic outer segment accompanied by increase of microtubule number and appearance of regularly spaced electron-dense bodies at tubular doublets; (f) branching of dendritic outer segment; and (g) transformation of the dendritic branches into curled lamellae and partial destruction of the dendrite sheath. The unique process of sensory cell degeneration is interpreted as an event that revokes a step towards a possible functional improvement of the labial palp-pit organ during further evolutionSupported by the Deutsche Forschungsgemeinschaft (SFB 4/G1)     

亚洲东部毛冠果蝇亚属修订（双翅目：果蝇科：冠果蝇属）

基于李彤等(2012)对冠果蝇属组(Stegana genous group)3个属间系统发育分析的结果,对亚洲东部冠果蝇属毛冠果蝇亚属(Orthostegana)进行了修订:将独冠果蝇亚属(Anastega)合并至毛冠果蝇亚属;记述了4新种:黄尾冠果蝇Stegana(O.)flavicaudua,密毛冠果蝇Stegana(O.)hirsutinna,林栖冠果蝇Stegana(O.)hylecoeta和多鬃冠果蝇Stegana(O.)multicaudua;补充描述了已知种弯脉冠果蝇Stegana(O.)curvinervis(Hendel,1914)和独冠果蝇Stegana(O.)singularis Sidorenko,1990;列表提供了上述6种毛冠果蝇间mtDNA COI & ND2基因序列的遗传距离。     

Evaluation of entomopathogenic nematodes against the olive fruit fly, Bactrocera oleae (Diptera: Tephritidae)

Infectivity of six entomopathogenic nematode (EPNs) species against Bactrocera oleae was compared. Similar infection levels were observed when third-instar larvae were exposed to infective juveniles (IJs) on a sand-potting soil substrate. When IJs were sprayed over naturally infested fallen olives, many larvae died within treated olives as well as in the soil; Steinernema feltiae caused the highest overall mortality of 67.9%. In addition, three laboratory experiments were conducted to optimize a time period for S. feltiae field application. (1) Abundance of fly larvae inside fallen olives was estimated over the 2006–2007 season with the highest number of susceptible larvae (3 mm and larger) per 100 olives being observed during December, 2006. (2) S. feltiae efficacy against fly larvae dropped to the soil post-IJ-application was determined. B. oleae added to the substrate before and after nematode application were infected at similar levels. (3) Effect of three temperature regimes (min–max: 10–27, 6–18, and 3–12 °C) corresponding to October through December in Davis, California on S. feltiae survival and infectivity was determined. After 8 weeks, the IJs at the 3–12 °C treatment showed the highest survival rate. However, the cold temperature significantly limited S. feltiae infectivity. Our results demonstrate that B. oleae mature larvae are susceptible to EPN infection both in the soil and within infested olives. Being the most effective species, S. feltiae may have the potential to suppress overwintering populations of B. oleae. We suggest that November is the optimal time for S. feltiae field application in Northern California.     

Natural establishment of a parasitoid complex on Bactrocera latifrons (Diptera: Tephritidae) in Hawaii

Bactrocera latifrons (Hendel) (Diptera: Tephritidae) is the most recent of four tephritid fruit fly species accidentally introduced into Hawaii. Although parasitoids have been released against other tephritid fruit fly species and have shown partial success in Hawaii, no parasitoids were released until 2004 to suppress populations of B. latifrons. The present study was conducted to document the parasitoid complex that has naturally established against B. latifrons in Hawaii and to assess whether there is a need for improving the biological control of this species. Based on ripe turkeyberry (Solanum torvum Sw) fruit collections over three consecutive years B. latifrons was the dominant tephritid fruit fly infestating turkeyberry at all four sites surveyed, across three major islands in Hawaii. The overall percentage parasitism of B. latifrons ranged from a low of 0.8% (Hana, Maui) to a high of 8.8% (Kahaluu, Oahu). Five primary parasitoid species were recovered from individually held B. latifrons puparia: Fopius arisanus (Sonan), Psyttalia incisi (Silvestri), Diachasmimorpha longicaudata (Ashmead), D. tryoni (Cameron), and Tetrastichus giffardianus Silvestri. F. arisanus was the predominant parasitoid at three of the four sites. Low levels of parasitism suggest that there is a need to improve biological control of B. latifrons, to minimize chances of this species causing economic impacts on crop production in Hawaii. We discuss the possibility of improving biological control of B. latifrons through augmentative releases of F. arisanus or introduction and release of specific and efficient new parasitoid species.     

Larviposition, Host Cues, and Planidial Behavior in the Sound-Locating Parasitoid Fly Homotrixa alleni (Diptera: Tachinidae)

Some groups of tachinid flies deposit mobile first-instar larvae (or planidia) on or near their host. Flies within one such group, the tribe Ormiini, parasitize singing species of ensiferan Orthoptera and use sound for long distance host location. However, what induces tachinids to larviposit and whether planidia use any cues actively to locate their host remains poorly known. This paper examines the larviposition and planidial behavior of the ormiine Homotrixa alleni in relation to its bushcricket host, Sciarasaga quadrata. Sound alone was sufficient to elicit larviposition in gravid female H. alleni, where females arriving at an arena placed over a speaker broadcasting host song deposited an equal number of planidia in the presence or absence of a silent S. quadrata. Flies were observed to larviposit by forcibly expelling planidia up to 6 cm in a forward direction from the fly, with less than half of the trials with a host present resulting in physical contact between the host and the fly. In the host's absence, flies walked around the arena significantly more often, remained on the arena for the experimental duration (10 min), and changed orientation frequently. In the host's presence, flies generally maintained a position facing the host, stayed in the quadrant of first approach, and typically flew off the arena within 2 min of arrival. When the oncoming fly approached a forward facing host, more planidia were found in the arena's center (i.e., closer to the host) than in the no-host or rearward-facing host treatment. Planidia experimentally placed on a circular arena averaged 1 cm of movement in 15 min but none of the following cues–host song, host song with song-vibration transmission, a silent host, and a silent host with host movement-vibration transmission–significantly affected the direction or distance planidia traveled. At 20 ± 1°C, over half of the planidia died within 1 h and all died within 2 h of deposition. The significance of these results in relation to reproductive strategies and parasitism is discussed.