Sensory interaction during trap-finding by female onion flies: implications for ovipositional host-plant finding

Using various three-dimensional traps alone and in combination with the onion volatile, dipropyl-disulphide (Pr₂S₂), we found that visual behaviour of female onion flies, Delia antiqua (Meigen), varied with the visual context (background composition and trap spacing) in which traps were presented and the females' reproductive state (mated vs. unmated). Against a background of real onions, females alighted more frequently on spherical than cylindrical traps, and white than green spheres, but females responded equally to white and green cylinders. In an onion field, baiting traps with Pr₂S₂ significantly increased female response to white over green traps, but had no influence on their response to trap shape. When traps were spaced 10 m apart and against a background of bare soil, females orienting to Pr₂S₂ baits alighted more frequently on vertical than horizontal traps, but shape and spectral reflectance were insignificant. However, when traps were spaced 0.25 m apart, females orienting to Pr₂S₂ baits alighted more frequently on cylinders than spheres. Mated females alighted more frequently on green than white cylinders, but unmated females responded to cylinders independent of spectral reflectance. When located 20 m downwind from Pr₂S₂ baits, mated females alighted on green cylinders significantly more often than unmated females. Response to traps mimicking onion plants suggests that ovipositional host-finding in female onion flies is dominated by olfactory responses at long range (several metres) and by visual cues at short-range (ca. 1 m). The view that host location by female onion flies is a hierarchical response pattern mediated by multiple sensory modalities and modified by resource level (habitat) and reproductive status, is discussed.     

Courtship behavior and discrimination between potential mates by maleDelia antiqua (Diptera: Anthomyiidae)

The repertoire of courtship behaviors of male onion maggots,Delia antiqua (Meigen), in a laboratory bioassay chamber, was analyzed by direct observation and by video recordings, in conjunction with a multichannel event recorder. Seven courtship behaviors were categorized: inspection from the substrate, aerial inspection, contact from the substrate, contact from the air, genital alignment, copulation, and male-male interaction. The frequency distribution of copulation bouts was best described by a Poisson distribution; peak mating activity occurred about 1 h into the bioassay. The duration in copulo, however, was extremely variable. On average, males spent 30 sin copulo (n=183); <30% of bouts were >50 s. The ability of males to discriminate between sexes, sexually immature and mature females, and between females ofD. antiqua and the cabbage maggot,Delia radicum (L.), was most pronounced in the elements of genital alignment and attempted copulation. The courtship and mating behavior inD. antiqua is consistent with a sequence that relies initially primarily on indiscriminate visual recognition of a potential mate, followed by species-and sex-specific semiochemical recognition upon contact.     

Mechanisms of resistance to onion fly egg-laying

When gravid onion fly females, Delia antiqua (Meigen) (Diptera: Anthomyiidae) were presented in laboratory choice test with 6-week-old plants of a susceptible cultivar and of onion breeding lines selected for resistance, mean numbers of eggs laid ranged from 34.8 to 1.6 eggs per plant. Differences in ovipositional responses were mirrored by differences in plant size. Analysis of covariance revealed no significant differences in ovipositional responses to breeding lines when differences in size were taken into account. Foliar surrogates were developed so that single size parameters could be varied while holding all other plant stimuli constant. Tests using these surrogates revealed that among plants with basal diameters of 1 to 4 mm and heights of 100 to 350 mm, diameter alone significantly influenced responses of ovipositing females. Ovipositional responses to plants beyond this size range could not be explained strictly by diameter differences. These results underscore the necessity of considering plant size as well as plant chemistry and texture in efforts to evalute and design resistant cultivars.

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Résumé A la lumière d'opinions récentes concernant la découverte des plantes-hôtes par D. antiqua, nous avons examiné les conditions de la résistance chez plusieurs lignées de A. cepa les moins attaquées lors d'essais en plein champ aux Pays-Bas, et dont on considérait que la résistance était due à une antixénose.Des femelles gravides en présence, au laboratoire, d'un choix de plants de 6 semaines, pondent en moyenne 1,6 à 34,8 oeufs par plant. Les différences entre les pontes correspondent aux tailles des plants. Si l'on tient compte de ces dernières, l'analyse de covariance ne montre pas de différence significative entre les pontes sur les différentes lignées. Des substituts de feuilles ont été créés de façon à faire varier les paramètres de taille tout en maintenant constants les autres stimuli de la plante: pour des plantes, dont le diamètre à la base variait de 1 à 4 mm et la hauteur de 100 à 350 mm, seul le diamètre a significativement influé sur la ponte. Les pontes sur les plantes hors de ces dimensions n'ont pu être expliquées par les seules différences de diamètre.Comme les paysans des principales régions productrices d'oignons d'Europe et des U.S.A. font des bénéfices importants avec la culture d'oignons précoces, ces lignées, sélectionnées par mégarde pour leur développement lent, ne sont probablement pas une bonne source de résistance à la mouche de l'oignon. Cependant, ces résultats soulignent la nécessité de tenir autant compte de la taille de la plante que de son chimisme ou de sa texture lors de l'évaluation et de la création de cultivars résistants.

     

Ovipositional Phenology and Behavior of the Spruce Cone Fly, Strobilomyia neanthracina (Diptera: Anthomyiidae)

The behavior of Strobilomyia neanthracina Michelsen, a phytophage infesting spruce (Picea spp.) seed cones, was observed at a field site in northern Ontario and in cages in a greenhouse to investigate spatiotemporal aspects of mating, host location, and oviposition. In the field, adults emerged from 21 to 24 May 1996, which coincided with bud burst of Picea glauca (Moench) Voss seed cones. For 4 days following emergence, Strobilomyia flies could no longer be seen at a monitored P. glauca tree and may have been on a mating or dispersal flight. Subsequently, females but not males were seen again and the oviposition period of ca. 3 weeks began. Mating was observed only in the greenhouse, mostly (i.e., 65%) at age 5–9 days. Although copulations lasted 11–45 min, these females laid infertile eggs only, beginning at age 4 days. No sperm was found in the spermathecal capsules of females, suggesting that no sperm had been transferred during these copulations. In both the field and the greenhouse, ovipositional sequences that resulted in egg deposition occurred throughout the day but few sequences were observed before 1000, probably because flies were not very active at air temperatures below 14°C (most sequences occurred at 25–27°C). In the greenhouse, the typical ovipositional sequence lasted an average of 7 min and consisted of landing on the cone and examining it with the proboscis and sometimes the ovipositor, egg deposition, and postovipositional behaviors such as tapping (touching the cone surface with the flabellum ca. 5 times s ^–1 ), which possibly represents a host marking behavior. In the field, tapping was seen less frequently than in the greenhouse but occurred significantly (P = 0.014) more often after sequences that resulted in egg deposition than after sequences that did not. Eggs hatched after 4–5 days at 20°C. In the greenhouse, the median longevity of females and males was 24 and 17 days, respectively.     

Identity and bioactivity of oviposition deterrents in pine oil for the onion maggot, Delia antiqua

The oviposition deterrent properties of pine oil (Norpine 65, Northwest Petrochemicals, Anacortes, Washington) for the onion maggot, Delia antiqua (Meigen), were verified using a two-choice bioassay with onion oil as an attractive control. The principal deterrent property of this pine oil was found to reside in three monoterpenes, 3-carene, limonene, and p-cymene, which were the primary constituents identified in the most deterrent of two fractions made by preparative gas chromatography of steam-distilled pine oil. At release rates of 220, 320 and 320 g per 24 h in two-choice bioassays these monoterpenes respectively caused 73.2, 65.4 and 56.3% deterrency of oviposition, while the ternary mixture released at 320 g per 24 h caused 88.6% deterrency. The ternary mixture also caused 62.5% deterrency in a no-choice bioassay. Of eight other monoterpenes tested, myrcene, -phellandrene, -terpinene, -phellandrene, -terpinene, terpinolene, and -pinene were significantly deterrent in declining order, while -pinene was inactive. The ternary mixture was released from glass capillary tubes or flexible plastic cylinders in further bioassays that challenged caged females to oviposit around the base of 35 potted onion seedlings with release devices placed on the soil surface. The most effective deterrency (85.3%) was achieved at a release rate of 280 g per 24 h per pot if plastic cylinder devices were deployed 24 h before the treated pot was exposed to D. antiqua females. If female D. antiqua were given only a treated pot, deterrency of oviposition on potted onion seedlings was significant, but low (11.7–63.2%). Because of incomplete efficacy, a monoterpene-based deterrent formulation would be best used operationally if combined with other deterrents, or if it were integrated with some other tactic.     

Pesticide-induced mortality of natural enemies of the onion maggot,Delia antiqua [Dip.: Anthomyiidae]

Pesticide-induced differential mortality betweenDelia antiqua(MEIGEN)and several other organisms associated with its natural control [D. platura(MEIGEN),Coenosia tigrina (F.),Entomophtora muscae (COHN), andAphaereta pallipes (SAY)] were evaluated under simulated field conditions. Direct and residual differential mortalities were described for 3 herbicides (Chloro-IPC, nitrofen, and CDAA), 3 fungicides (maneb, chlorothalonil, and copper sulfate), and 1 insecticide (malathion). The recommended field application rates of these chemicals produced high positive differential mortality levels betweenD. antiqua and some of the natural mortality agents. Chloro-IPC, a preplant and midseason herbicide, induced 100% mortality ofA. pallipes over the 3-day residual test period. Modifications in behavioral orientation and death patterns ofD. antiqua adults infected withE. muscae were also noted in malathion-treated flies such that subsequent conidial dispersal was highly restricted. Michigan Agricultural Experimental Station Journal Article Number 10133.     

四川省花蝇科5种雌性的记述(双翅目:蝇总科)

分别记述采自四川西部山区的小片花蝇(Anthomyia parvilamina Feng,1987)、棘基植种蝇[Botanophila spinulibasis(Liet Deng,1981)]、离叉植种蝇[Botanophila apodicra(Feng,1987)]、缘齿华种蝇(Sinohylemya craspedodonta Hsue,1980)、中华泉蝇(Pegomya chinensis Hennig,1973)等5种花蝇科雌性的模式产地及外形特征。     

Choices and Consequences of Oviposition by Hylemya (Delia) Sp. (Diptera: Anthomyiidae)

Hylema sp.⁵females oviposit on the undersides of sepals of developing buds of both Ipomopsis aggregata and Polemonium foliosissimum. Eggs deposited on the latter are significantly more likely to be fully protected by the sepal than are eggs deposited on the former. Unexposed eggs have a significantly greater likelihood of successfully developing to the larval stage than do exposed eggs. The difference in frequency of egg exposure on the two plant species can be attributed to differences in sepal morphology: I. aggregata sepals are significantly narrower than those of P. foliosissimum. The hypothesis that females preferentially oviposit on larger flowers was unconfirmed by a manipulative choice experiment. Plants differing in the size of their flowers were potted together and presented to Hylemya in arrays in the field. Flowers of the larger-flowered pair were no more likely to be oviposited on than flowers of the smaller-flowered pair. However, there were significant negative correlations between the corolla length and the percentage of flowers laid on per day at each of two sites. There was also a significant positive correlation between the corolla width and the percentage of flowers laid on at one site. Thus females appear to be using some measure of flower morphology, or a correlated trait, in making oviposition decisions. The degree to which Hylemya is making suboptimal choices between host plant species is discussed and requires further examination.     

Fitness of a translocation homozygote in cage experiments with the onion fly,Hylemya antiqua (meigen)

Summary In Hylemya antiqua, a stock homozygous for an autosomal reciprocal translocation was isolated using egg-hatch reduction and karyotype analysis. Sibling translocation homozygous (TT) and heterozygous (T+) females were compared in respect to egg production and longevity. In one full-sib (5 TT and 8 T+ females) significantly higher values for both parameters for T+ than for TT females were scored, in four others (a total of 35 TT and 28 T+ females) no significant differences were found. Cage experiments were started with populations composed of equal numbers of wild type flies (++) and translocation homozygotes. The frequencies of the different karyotypes in three successive, non-overlapping generations, did not suggest substantial differences in fitness between ++ and TT flies. Possible causes of a surplus of T+ individuals found in these experiments are discussed together with the usefulness of the translocation for genetic control of H. antiqua.     

Physical and chemical factors influencing landing and oviposition by the cabbage root fly on host-plant models

Various plant models were used in both choice and no-choice tests in the laboratory to assess landing and oviposition preferences of the cabbage root fly,Delia radicum (L.). The main factor governing the site most suitable for landing was the conspicuousness of the object and not its shape. Oviposition was influenced considerably by the pre-conditioning of the females. Deprived females laid eggs even when denied access to both host plant chemicals and host-plant models. The dominant role of contact chemical stimuli in host acceptance was reconfirmed, but only a combination of physical and chemical stimuli appeared capable of eliciting normal oviposition. The combination of contact chemical stimuli and the presence of a stem on the test model had a synergistic effect on the numbers of eggs laid in both choice and no-choice situations. In choice bioassays, female cabbage root flies distinguished between models of different shapes, heights and sizes. The size and shape of the models appeared to be perceived in part after the flies had landed.     

Effects of the angle of inclination of traps on the numbers of large Diptera caught on sticky boards in certain vegetable crops

The angle of inclination of the surface on which flies prefer to land in vegetable crops was studied in field plots and in field-cages using one-sided sticky traps aligned in one plane but orientated in eight directions. The four Delia species studied, D. antiqua, D. floralis, D. platura and D. radicum, preferred to land on horizontal surfaces, indicating that they are likely to be trapped in largest numbers on traps with a horizontal trapping surface, such as water traps. This trend was even more pronounced with the Syrphidae. In contrast, greatest numbers of the carrot fly, Psila rosae, were caught on the lower surface of traps inclined at 45° to the vertical. The advantages of using traps inclined in this way for trapping P. rosae are that more flies are caught on such surfaces, the sticky trapping compound is protected from the adverse effects of rain, and the traps are highly selective.

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Résu,é L'angle d'inclinaison de la surface sur laquelle les mouches préfèrent atterrir dans les cultures de légumes a été examiné dans la nature, dans des parcelles et dans des cages, en utilisant des pièges englués sur un côté, alignés dans le même plan, mais orientés dans 8 directions. Les 4 espèces de Delia étudiées: D. antiqua, D. floralis, D. platura, D. radicum, ont préféré atterrir sur des surfaces horizontales, ce qui implique qu'elles sont probablement prises en plus grand nombre sur des pièges avec une surface de capture horizontale, comme des pièges liquides. Cette tendance est encore plus prononcée chez les Syrphidae. A l'opposé, de nombreuses mouches de la carotte, Psila rosae, ont été capturées sur la face inférieure de pièges inclinés de 45°. Il est avantageux d'utiliser des pièges inclinés de cette façon contre P. rosae parce que, d'une part ils sont très efficaces, d'autre part le produit actif est protégé de la pluie et enfin, le piège est très sélectif.

     

Influence of background on host-plant selection and subsequent oviposition by the cabbage root fly (Delia radicum)

The behaviour of female cabbage root flies during host plant selection was studied in the laboratory using brassica plants growing in backgrounds of bare soil, clover, grass, peas and four non-living materials. Gravid females landed about twice as often on brassica plants growing in bare soil than on comparable plants growing amongst non-host plants. Once a receptive female landed on a brassica plant, the female made, on average, four ‘spiral flights’ and two jumps on and off the plant before laying alongside the plant. Surrounding a brassica plant with a diverse background altered the behaviour of the flies, so that the spiral flights around the host plant were replaced by short hops between nearby vertical objects. The loss of contact and recontact with the host plant then prevented the females from accumulating sufficient contacts with the host plant to be stimulated to lay. Spiral flights around host plants appear to determine whether or not flies will lay alongside host plants. Flies in mixed plantings have a reduced rate of settling on the host plant, and a higher rate of locomotion, because they land frequently on non-host plants. Hence, visual stimuli appeared to have greater effects than, chemical or mechanical barriers in deterring flies from laying alongside brassica plants in diverse backgrounds. In ‘choice’ situations, backgrounds of real plants reduced oviposition alongside brassica plants by at least 50%. In ‘no-choice’ situations, flies laid similar numbers of eggs alongside all brassica plants irrespective of plant background or plant size. If numbers of fly eggs are to be reduced on commercial brassica crops by undersowing the crops with clover, plants growing in bare soil may also have to be included to provide the flies with sites preferred for oviposition.     

Laboratory transmission and in vivo incubation of Entomophthora muscae (Entomophthorales: Entomophthoracae) in the onion fly,Delia antiqua (Diptera: Anthomyiidae)

A high-moisture infection chamber was used for the in vivo transmission of Entomophthora muscae within laboratory populations of the onion fly, Delia antiqua. This cadaver-to-fly transmission procedure provided an average experimental infection rate well above 95%. Laboratory infection and temperature-dependent incubation rates of E. muscae were further examined in adult populations of D. antiqua. The time from initial exposure until host death and pathogen sporulation was accurately predicted using a second-order function of the incubation temperature. A developmental base temperature of approximately 5°C was estimated, with 105 degree-days being the average number of heat units required between host infection and death. E. muscae transmission between D. antiqua and D. platura, two insect pests typically associated with Michigan onion production, was verified under laboratory conditions.     

中国四川花蝇科三新种及一新记录种(双翅目:蝇总科)

报道采自中国四川西部花蝇科(Anthomyiidae)3新种:隰蝇属的乌亮隰蝇(Hydrophoria nigrinitidaFeng,sp.nov.);壮叶隰蝇(H.robustisurstylusFeng,sp.nov.);泉蝇属的黑前足泉蝇(Pegomya nigripraepedaFeng,sp.nov.)。模式标本存在中国科学院上海生命科学研究院植物生理生态研究所。同时记录泉蝇属(PegomyaRob ineau-Desvoidy,1830)的一中国新记录种:丽足泉蝇[Pegomya pulchripes(Loew,1857)]。     

Meiotic disjunction and embryonic lethality in sex-linked double-translocation heterozygous males of the onion fly,Hylemya antiqua (Meigen)

Summary The frequencies of disjunction types in double-translocation heterozygous males (2⁶2^Y6²6XY²) in Hylemya antiqua have been established in MII cells and eggs of testcrosses.Several disjunction types occurred but four predominated. A correlation was found between the frequencies of the disjunction types and the relative position of the centromeres. The frequency of numerical non-disjunction (NND) was 4%. Differences in frequency of NND between sex-linked and autosomal translocations of H. antiqua are discussed. A good correspondence between the frequencies of unbalanced karyotypes, and embryonic and larval mortality was found. The total genetic load which can be induced by the T14/T61 males is estimated to be 60–65%. Some duplication/deficiency karyotypes appeared to be viable in pupal and even adult stages. In 2⁶2⁶2^Y6²6²X males a regular coorientation between 2^Y and X was observed, in spite of non-homologous centromeres and a complicated synapsis of 2^Y. Application possibilities of the present material for genetic control of H. antiqua are discussed.     

Completion of diapause in field populations of the cabbage root fly (Delia radicum)

The various diapause and post-diapause stages entered by cabbage root fly pupae during the overwintering period are shown schematically. Although diapause induction started in mid-Aug., the early-pupating insects did not develop further but were maintained in diapause by the warm autumn temperatures. Therefore, diapause development was simultaneous in all Wellesbourne pupae, whether of second or third generation origin. Diapause development started only in mid-Oct., when mean soil temperatures fell below 10°. In the field, 90% of the overwintering population of cabbage root fly pupae had completed pleted diapause by 5 March 1980, 17 Feb. 1981 and 18 Feb. 1982. This was equivalent to a duration of 19 weeks from mid-Oct. onwards, during the winters of 1979–80, 1980–81 and 1981–82 respectively. A further break between the completion of diapause and the warm conditions required to start post-diapause development also helps to condense the emergence of flies in the spring. Hence, an accurate forecast of the time of spring attack by populations of flies similar to those at Wellesbourne should be possible.This study was financed partly by the Commission of the European Communities as CEC Contract No. 0771.     

Visual behaviour of the onion fly, Delia antiqua: antagonistic interaction of ultraviolet and visible wavelength reflectance

The interaction of ultraviolet (UV=350–400 nm) and visible (VISIBLE=400–650 nm) wavelengths in the visual behaviour of the onion fly, Delia antiqua (Meigen), was examined by measuring spontaneous alightment on various UV- and non-UV-reflecting, sticky cardboard traps in onion fields. Alightment on traps was negatively correlated with the percent UV (350 nm) and green (540–580 nm) reflectance and positively correlated with the percent blue (430–470 nm) reflectance. Alightment varied directly with the ratio of stimulatory/inhibitory reflected wavelengths. Males and females were similar in their response to both UV and VISIBLE wavelength reflectance, with the exception that males were more sensitive than females to UV-reflecting white surfaces. A multiple regression model, that used the intensity of 3 key wavelengths, 350, 450 and 560 nm, as independent variables, explained 90% of the variation in the combined male and female response to spectral reflectance from traps. These results indicate that some visual behaviours of D. antiqua are a function of the integration of sensory input from the entire spectral distribution of the stimulus, and not simply the dominant wavelengths or hue.

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Résumé L'interaction des ondes ultraviolettes (UV=350–400 n nm) et des ondes visibles (VISIBLE=400–650 nm) sur le fonctionnement visuel de la mouche de l'oignon, Delia antiqua, a été examinée par le mesurage des descentes spontanées sur des pièges variés, construits de carton gluant, qui reflète le UV ou non, dans des champs d'oignons. Les descentes furent en rapport négativement avec le pour-cent de la réflexion de l'UV (350 nm) et du vert (540–580 nm), et en rapport positivement avec la réflexion du bleu (430–470 nm). Le chiffre des descentes était en rapport directement avec la porportion stimulantes/inhibitantes des ondes reflétées. Les mâles et les femelles étaent comparables en leurs réponses aux ondes UV et VISIBLES reflétées, sauf que les mâles fussent plus sensibles que les femelles aux surfaces blanches et UV-reflétantes. Une modèle d'analyse multiple de regression qui prenait en considération les intensités des trois ondes significatives, 350, 450 et 560 nm, en tant que les variables indépendants, a expliqué 90% de la variation en la réponse combinée des mâles et des femelles à la réflexion spectrale des pièges. Ces resultats indiquent que quelques fonctionnements visuels de la D. antiqua sont une fonction des données sensorielles de la distribution spectrale entière du stimulus, pas seulement une fonction des ondes dominantes ou de la teinte.

     

Oviposition behavior of an ant-parasitizing fly,Neodohrniphora curvinervis (Diptera: Phoridae), and defense behavior by its leaf-cutting ant hostAtta cephalotes (Hymenoptera: Formicidae)

This study examines the oviposition behavior of the phorid parasitoid Neodohrniphora curvinervisand the antiparasitoid defense behavior of its leafcutting ant host Atta cephalotes. N. curvinervisfemales are diurnal sit- and- wait parasitoids that attack only outbound foragers of head width 1.6 mm or greater. Females deposit a single egg through the foramen magnum of each host successfully parasitized. Pursuit of hosts is usually initiated when an outbound forager of acceptable size passes by a parasitoid perch site. Individual foragers defend themselves against pursuing parasitoids by outrunning them along the foraging trail or by standing their ground and fending them off with their legs,antennae, and mandibles. At the colony level, susceptible foragers are protected against parasitism by a shift in the forager size distribution toward smaller unsusceptible sizes during the day when parasitoids are active and toward larger sizes at night when parasitoids are inactive. The frequency of parasitism of susceptible foragers was 15%, which is more than five times the frequency found in another system involving the phorid parasitoid Apocephalus attophilusand the leafcutting ant host Atta colombica.We offer several possible explanations for such differences in the frequency of parasitism and also examine reasons for the high incidence of superparasitism (19%) observed in the system studied.     

TheMoegistorhynchus longirostris (Diptera: Nemestrinidae) pollination guild: long-tubed flowers and a specialized long-proboscid fly pollination system in southern Africa

A guild of 20 late spring- and early summer-flowering species ofIridaceae, Geraniaceae andOrchidaceae is pollinated partly or exclusively by the long-proboscid flyMoegistorhynchus longirostris (Nemestrinidae). This large-bodied fly, active in late spring and early summer, has mouthparts 40–70 (90) mm long and forages for nectar from a variety of species. These plants share a suite of convergent floral features including a straight or weakly curved floral tube usually 50–70 mm long but sometimes to 90 mm, relatively short petals or tepal lobes coloured white, cream or salmon with reddish nectar guides, and often violet or red anthers and pollen. Flowers of most species with these characteristics are zygomorphic with the stamens either arcuate (mostIridaceae) or declinate (Geraniaceae and someIridaceae). The flowers are odourless and typically secrete large amounts of nectar of relatively constant sugar concentration, mostly 24–29%, with a high sucrose:hexose ratio. Guild members utilize five separate sites of pollen deposition on the body of the fly, typically utilizing different deposition sites when two or more co-occur, indicating strong selection to aviod pollen contamination.M. longirostris is restricted to the west coast of southern Africa and at least 8 species appear to depend exclusively on the insect for pollination. The remaining species in the guild are pollinated by one or both of the long-proboscid fliesPhiloliche gulosa andP. rostrata (Tabanidae) over other parts of their range. Species and races pollinated entirely byM. longirostris have longer floral tubes which makes nectar unavailable to other insects, including other species of long-proboscid fly. The only insect with mouthparts long enough to forage effectively on these long-tubed flowers isM. longirostris and this fly must be considered a keystone species in the ecosystems in which it occurs.Dedicated to emer. Univ.-Prof. DrFriedrich Ehrendorfer on the occasion of his 70th birthday