The use of common elder Sambucus nigra to promote Aphidophagous syrphids in apple orchards

Elder shrubs (Sambucus nigra L.) were planted in an experimental apple orchard as bordering hedgerow with the objective of rearing syrphids (Diptera: Syrphidae) and consequently enhancing the biological control of the rosy apple aphid Dysaphis plantaginea (Passerini). The inoculative introduction of the specific elder aphid Aphis sambuci L. (Homoptera: Aphididae) in late March 2002 and again in 2003 led to the complete hedgerow turning into an early and productive alternative prey reservoir for indigenous syrphids. The species Scaeva pyrastri (L.), S. selenetica (Meigen), Syrphus ribesii (L.), S. vitripennis Meigen, Epistrophe eligans (Harris), E. nitidicollis (Meigen), Platycheirus scutatus (Meigen), Eupeodes corollae (F.), Meligramma triangulifera (Zetterstedt) and Episyrphus balteatus (De Geer) developed on the elder hedgerow during the spring 2002 taking advantage of the aphid infestation. The season 2003 was characterised by a quite different species abundance pattern as, contrary to the previous year when both monovoltine and polyvoltine species were equally represented, the monovoltine syrphids (Epistrophe spp.) represented the dominant group exploiting the elder aphids. A parallel analysis showed that the faeces of the first adult syrphids observed ovipositing on the elder hedgerow in spring contained digested pollen of mainly apple. Our observations indicate the possibility of establishing a local population of monovoltine syrphids in apple orchards by managing an aphid-infested elder hedgerow, without any additional pollen or nectar producing plants.     

Why are there good and poor mimics?

Among the many Batesian mimetic hoverflies (Diptera: Syrphidae) some have a very precise resemblance to the presumed model ('good' or 'specific' mimics) while others have a much less precise resemblance ('poor' or 'general' mimics). Intuitively one might expect that the specific mimics would be commoner and more successful than the general mimics. However, many specific mimics (e.g. Serkomyia silentis, Volucella bombylans) are quite rare, while general mimics are common (e.g. Syrphus ribesii, Episyrphus balteatus and Eristalis intricarius). Similarly, some ant-mimicking spiders from several different families are very good morphological and behavioural mimics of just one species of ant while others have a less detailed resemblance to ants in general. Six hypotheses are presented to explain the occurrence of so many poor mimics, and a theoretical model is outlined (the multi-model hypothesis) which shows how a poor general mimic can have a larger population than a good, specific mimic. This hypothesis may apply to some species of hoverfly and to some ant-mimicking spiders.     

Hoverflies of the Guelma district,with species new to Algeria and North Africa (Diptera: Syrphidae)

A survey of the hoverflies (Diptera: Syrphidae) of the region of Guelma, northeast Algeria, was carried out from August 2011 to July 2013. A total of 31 species was recorded with six (Scaeva mecogramma, Epistrophe melanostoma, Syrphus vitripennis, Brachypalpus valgus, Merodon calcaratus and M. chalybeus) new to Algeria. Of these, three species (E. melanostoma, M. calcaratus and M. chalybeus) are also new additions to the entomofauna of North Africa. The results complement those of a previous study covering Numidia and Tebessa and provide a more comprehensive view of the phenology and distribution of Syrphidae across northeast Algeria. An improved knowledge of the distribution of North African hoverflies will contribute to a better assessment of the status of each taxon and inform conservation efforts.     

Predation and potential transfer of pollen in a population of Saxifraga hirculus

Body surface pollen load and gut pollen (pollen diet load) of flower visitors in a Danish population of Saxifraga hirculus were analyzed. The four most frequent visitors, Eurimyia lineata (Syrphidae), Neoascia tenur (Syrphidae), Asindulum nigrum (Mycetophilidae), and Zygaena trifolii (Zygaenidae) had total surface loads (and S. hirculus loads) of 1031 (771), 137 (106), 108 (78), and 318 (27) pollen grains, respectively. Other syrphid visitors had on average a load of 407 (192) pollen grains. Males of E. lineata, N. tenur , and A. nigrum carried more surface pollen than the females; females of E. lineata had a lot of pollen from Ranunculus acris and S. hirculus and males had a lot from S. hirculus . Females of N. tenur carried pollen mostly from S. hirculus, Lotus uliginosus and Galium uliginosum ; males carried pollen mostly from S. hirculus . The surface pollen load of A. nigrum was dominated by S. hirculus pollen, whereas that of Z. trifolii was dominated by Cirsium palustre and Lychnis flos-cuculi pollen. The amount and kind of pollen varied enormously both intraspecifically and interspecifically. Eurimyia lineata, N. tenur and other syrphids had total pollen diet loads (and S. hirculus pollen diet loads) of approximately 14900 (657), 3739 (2327) and 17784 (13040). The average syrphid contained pollen from five plant species, only one to two, however, were quantitatively important. The average diet load in females was nine times that in males in both E. lineata and N. tenur. Eurimyia lineata ate a lot of R. acris; N. tenur ate a lot of S. hirculus and G. uliginosum . Generally, no correlation was observed between size or season and the composition of surface load and diet load. Eurimyia lineata made most of the pollen visits. Thus, the seed set from this Danish population of S. hirculus was dependent on a single syrphid species.     

Purification and characterization of glutathione S-transferases from two syrphid flies (Syrphus ribesii and Myathropa florae)

Glutathione S-transferases (GST) play an important role in the detoxification of many substances including organic pollutants and plant secondary metabolites. We compared the GST of two syrphid species, the aphidophagous Syrphus ribesii and the saprophagous Myathropa florea to assess the relation between feeding type and GST patterns. Differences between the GST of the hoverfly species were observed after purification by affinity chromatography, SDS-PAGE and kinetic studies. While the specific activities of the purified enzymes were different, the purification yields were similar. The variation in specific activities was related to the presence of different isoenzymes in both syrphid species by SDS-PAGE. While two bands of 24 and 32 kDa were observed for M. florea, one more band of 26 kDa was present in S. ribesii. When a range of substrate and glutathione concentrations was tested, differences in Km and Vmax between the glutathione S-transferases from both hoverfly species were also observed. These results are discussed in terms of adaptations to the feeding habit and the habitat of the two syrphid species.     

Hoverfly (Diptera, Syrphidae) fauna of Ary-Mas locality (the Taimyr Biosphere Reserve)

The local fauna of Ary-Mas (the Taimyr Biosphere Reserve) was found to comprise 56 species of hoverflies, representing only two subfamilies: Syrphinae (36 species) and Eristalinae (20 species). Analysis of trophic preferences of adult hover-flies showed that Syrphidae visited plants of 17 species from 10 families. The representatives of Syrphidae preferred Tanacetum bipinnatum (18 species), Ranunculus propinquus, and Tripleurospermum hookeri (12 species each). Three species of hoverflies visited the greatest number of plant species: Melangyna arctica (11), Syrphus ribesii (9), and Parasyrphus dryadis (8).     

The mesmerizing wart: the pollination strategy of epiphytic lady slipper orchid Paphiopedilum villosum (Lindl.) Stein (Orchidaceae)

Paphiopedilum uillosum was studied in hill evergreen forests (at up to 35 m above ground level) in North Thailand between 1990 and 1994. Flowering lasted 4.5 months: flower longevity 2–3 months. During 224 hours of flower-watching, less than 100 specimens of Syrphidae (Diptera) were seen in the vicinity of the flowers; 15 cases of pollen acquisition were mainly by female Eplsyrphus alternans, Syrphus fulvifacies, Betasyrphus serarius among 6 spp. of pollinators. Populations peaked during the main flower opening period, one month before the maximum number of open flowers was reached in early February. The flowers are 'kettle-traps' without known reward, luring mainly by food deception. Long distance attraction is probably by a urine-like odour (attraction to mammalian excretions had hitherto been little reported) and colour contrast. Close range lure is by the glittering staminode (probably mimicking droplets of honeydew/moisture) centred by a slippery wart (faking a perch) at which pollinators fly, immediately losing their grip and tumbling into the pouch, preventing wing action and falling being disrupted by a trough-shaped frame. Escape is up the tunnel, past the stigma, to the exit where, pressed by several mechanisms against the anther, the hoverfly scoops off very sticky pollen which has a viability of at least 8 weeks. Capsule formation averaged 8%.     

Pollen transport differs among bees and flies in a human‐modified landscape

Aim Dispersal distances of insect pollinators are critical in defining their contribution to landscape‐wide pollen movement and ultimately gene flow in natural and agricultural systems. We ask whether bee and fly pollinator taxa differ in their dispersal distances and transport of viable pollen in a human‐modified system. Location Canterbury and Otago region, South Island, New Zealand. Methods We captured pollen‐carrying insects travelling outside of a model mass‐flowering agricultural crop, Brassica rapa, using insect flight intercept traps at five distances (0, 100, 200, 300 and 400 m) from the pollen source. We examined pollen loads and pollen viability to determine whether pollen transport distance and viability differ among pollinator taxa. Results A total of 5453 insects were collected of which 717 individuals from 26 insect taxa were positively identified as dispersing pollen up to 400 m from the source. These taxa consisted of four species from two bee families (Hymenoptera: Apidae and Halictidae), and eight species from four fly families (Diptera: Bibiondae, Stratiomyidae, Syrphidae and Tachinidae). Apidae generally carried higher pollen loads and more viable pollen than most fly taxa. Taxa in the fly families Stratiomyidae and Syrphidae, however, carried pollen to 400 m, which is further than both bee families. Main conclusions A diverse array of wild and managed flower visitors can transport viable pollen from a pollen source to at least 400 m. Knowledge of the differences in transport distances among generalist pollinators in human‐modified environments is crucial to understand the potential extent to which (1) pollen transport can facilitate gene flow and (2) unwanted hybridization may occur between crops and related weeds.     

Oviposition preferences of aphidophagous hoverflies

1. Oviposition preferences of two predatory hoverflies, Episyrphus balteatus and Syrphus ribesii (Diptera, Syrphidae), were studied in the laboratory. 2. There was broad agreement between the two species: two of the top three preferred prey for oviposition in both hoverflies were pea and rose aphids; nettle aphids were consistently the least preferred. 3. Discrimination decreased with age.     

A photographic account of pollen digestion by adult hoverflies

ABSTRACT. The method of pollen digestion by adults of the hoverfly Cheilosia albitarsis Mg. (Diptera: Syrphidae) was investigated by observing pollen grains at different stages in their progression along the gut. No form of mechanical 'grinding' to break up the grains was observed, as suggested by some previous authors. Instead, there was an exudation of the pollen grain contents through one of the pores in the pollen exine. This exudation may start as a growing pollen tube. The exact conditions which cause the pollen grains to give up their contents in this way are unknown, but it is suggested that nectar sugar within the gut may provide the necessary stimulus.     

Comparison of the pollen content on the body and in the gut of hoverflies (Diptera,Syrphidae)

We analyzed the pollen content in the gut and on the body of hoverflies (Diptera: Syrphidae), mostly from three genera (Eristalis, Syrphus s. l., and Sphaerophoria) that visited flowers of two jointly flowering species of Apiaceae with indistinguishable pollen (Anthriscus sylvestris and Aegopodium podagraria) and of Bunias orientalis (Brassicaceae). Pollen compositions depended on the plant from which the flies were collected rather than on the insect genus. At the same time, individual preferences of the studied flies to food plant species were confirmed. The pollen compositions in the gut and on the body were weakly correlated. Two possible reasons for such a weak correlation are discussed: individual differences in the fly feeding habits and different time of pollen preservation in the gut and on the body.     

Pollen‐feeding in hover‐flies (Diptera: Syrphidae)

Analyses of the pollen contents of the crop and intestine of 11 species of New Zealand Syrphidae . showed that small, sparsely haired hover‐flies with unbranched hairs, short, simple bristles, and a short proboscis had ingested at least 99% anemophilous pollens, and that larger, more hairy hover‐flies with pollen‐collecting hairs, long, spirally grooved bristles, and elongate mouthparts had ingested pollens almost exclusively from nectar‐bearing flowers. Pollen‐feeding behaviour was studied in one hairy species, the drone‐fly Eristalis tenax, and in one sparsely‐haired species, Melanostoma fasciatum. Using granulated charcoal as a substitute for pollen, it was found that in E. tenax particles trapped among the body hairs are combed off by the front and hind tibiae and transferred to pollen‐retaining bristles on the front and hind tarsi respectively. Particles retained among the front tarsal bristles are ingested directly from the bristles. Those retained by the hind tarsi are transferred in flight by leg‐scraping movements to the front tarsi, from which they are subsequently eaten. E. tenax also eats pollen directly from anthers. In M. fasciatum apparently all the pollen ingested is taken directly from anther lobes or stigmas. The few pollen grains that adhere to the body of this species are combed off by the front and hind tibiae and transferred to the front and hind tarsi, but are not retained there because the bristles are short and simple. The mouthparts, hairs, and bristles of E. tenax and M. fasciatum are illustrated. Drawings of leg movements associated with pollen collection and ingestion, and photographs showing leg scraping in E. tenax are included. Morphological similarities between drone‐flies and honey‐bees, previously regarded as the result of mimicry, can be explained by convergent evolution in response to similar food‐gathering behaviour. Probably the majority of Syrphidae, and also the related Acroceridae, collect pollen by means of branched or curly‐tipped hairs.     

Factors affecting the freeze tolerance of the hoverfly Syrphus ribesii (Diptera: syrphidae)

Larvae of Syrphus ribesii collected from overwintering sites in the U.K. are strongly freeze tolerant with 70% survival at -35 degrees C. The cold tolerance of laboratory reared insects increased with increasing periods of acclimation at 0 degrees C, with a concurrent rise in the supercooling point (SCP) from -6.8+/-0.1 to -5.1+/-0.3 degrees C. There was 50% survival in the most cold-hardy group 72h after brief exposures to -30 degrees C. The retention of gut contents caused a decrease in cold hardiness, with only 13% of larvae surviving 72h after exposure to -15 degrees C, with no subsequent pupation or emergence. Wet larvae had a significantly higher SCP (-5.0+/-0.2 degrees C) compared to dry larvae (-7.8+/-0.4 degrees C), although survival of larvae was similar in both groups. There was no nucleator activity in the haemolymph of field collected larvae. The importance of these findings are discussed in relation to the freeze tolerance strategy of S. ribesii.     

Flowering and seasonal changes in flower sex ratio and frequency of flower visitors in a population of Saxifraga hirculus

The open, dish-shaped flowers of Saxifraga hirculus reflected ultraviolet and yellow light, contained very small amounts of nectar, and contained an average of about 75300 pollen grains per flower. Almost 11% of the pollen was inviable. Stigmatic pollen loads and seed set decreased during the course of the season. The plant appeared to be fully between-ramet compatible and partially within-ramet compatible. Seed set for the population was 30.3%. The protandrous flower opened during the day and had male and female phases of nine and three days, respectively. The protandrous system reduced the number of pollination days by a third.
At least 26 species of insects, 16 of which were syrphids, visited the flowers. Based on the number of flower visits, four species were the dominant visitors of S. hirculus: Eurimyia lineata and Neoascia tenur (Diptera: Syrphidae); Asindulum nigrum (Diptera: Mycetophilidae), and Zygaena trlfolii (Lepidoptera: Zygaenidae). Eurimyia lineata was the most frequent visitor (51% of all visits). As the season advanced, the visits by E. lineata decreased, whereas the visits by A. nigrum increased. Z. trifolli disappeared completely towards the end of the season. Only two thirds of the pollination days were "good" foraging days for these visitor species. The four major visitor species spent an average of 11.7, 27.4, 30.7 and 22.6 s per flower, respectively. Estimates suggest that about 6.5 visits (which is equal to 2.6 min of flower-visiting) and 2100 grains of pollen were required to produce one seed.     

Cover Caption

An adult flower fly (Syrphidae) rests on a flower of Erigeron annuus (Asteraceae) growing on a rice farm in eastern China. Plants of non‐crop species can be important in providing nectar and pollen to natural enemies of crop pests. Whilst the larvae of flower flies prey on pests such as aphids, the adults are non‐predacious and females need pollen in order to produce eggs. See pages 1–12. Photo by Ping‐Yang Zhu.     

A survey of syrphid predators of Nasonovia ribisnigri in organic lettuce on the central coast of California

Organic lettuce, Lactuca sativa L., producers on California's Central Coast rely on endemic syrphid flies (Diptera: Syrphidae) to suppress populations of Nasonovia ribisnigri Mosley (Homoptera: Aphididae) and other aphids affecting lettuce. Growers are using various forms of habitat manipulation to enhance biological control. We surveyed syrphids collected from organic romaine in and around the Salinas Valley from March through September 2005 to gain a better understanding of the species responsible for aphid suppression and to examine possible implications for biocontrol. The primary species of syrphid fly reared were Toxomerus marginatus (Say) (39%), Platycheirus stegnus (Say) (27%), Sphaerophoria sulfuripes (Thomson) (13%), and Allograpta obliqua (Say) (10%). Syrphus opinator Osten Sacken (2%), Toxomerus occidentalis (Curran) (1.3%), and Eupeodes volucris Osten Sacken (1%) were less common. Sphaerophoria pyrrhina Bigot, Scaeva pyrastri (L.), Platycheirus obscurus Say, Allograpta exotica Wiedemann, and Eupeodes americanus Wiedemann each made up <1% of the syrphids reared. T. marginatus and Sphaerophoria sulfuripes were commonly collected from romaine plants with few or no detectable aphids. P. stegnus was observed to deposit in clusters of eggs, and was only reared in significant numbers from highly infested fields. Approximately 5% of syrphid larvae overall were parasitized by either Diplazon sp. (Hymenoptera: Ichneumonidae) or Pachyneuron sp. (Hymenoptera: Pteromalidae).     

Convergence of chemical mimicry in a guild of aphid predators

Abstract.  1. A variety of insects prey on honeydew-producing Homoptera and many do so even in the presence of ants that tend, and endeavour to protect, these trophobionts from natural enemies. Few studies have explored the semiochemical mechanisms by which these predators circumvent attack by otherwise aggressive ants.
2. Ants use specific mixtures of cuticular hydrocarbons (CHCs) as recognition labels, but this simple mechanism is frequently circumvented by nest parasites that engage in 'chemical mimicry' of their host ants by producing or acquiring a critical suite of these CHCs.
3. Analysis of the CHCs from the North American woolly alder aphid, Prociphilus tessellatus (Homoptera: Aphididae), their tending ants, and aphid predators from three insect orders, Feniseca tarquinius (Lepidoptera: Lycaenidae), Chrysopa slossonae (Neuroptera: Chrysopidae), and Syrphus ribesii (Diptera: Syrphidae), showed that while the CHC profile of each predatory species was distinct, each was chemically more similar to the aphids than to either tending ant species. Further, the CHCs of each predator species were a subset of the compounds found in the aphids' profile.
4. These results implicate CHCs as a recognition cue used by ants to discriminate trophobionts from potential prey and a probable mechanism by which trophobiont predators circumvent detection by aphids and their tending ants.
5. Although several features of the aphids' CHC profile are shared among the chemically mimetic taxa, variation in the precision of mimicry among the members of this predatory guild demonstrates that a chemical mimic need not replicate every feature of its model.     

The phenology of Syrphidae (Diptera): are they Batesian mimics of Hymenoptera?

The phenology of aculeate Hymenoptera and of syrphids which are believed to mimic them has been investigated at three semi-natural ancient woodland sites in north-west England. It is concluded that the abundance and phenology of most of the hoverflies is consistent with their being Batesian mimics of particular species of bee or wasp. The main exceptions are Eristalis spp., Helophilus spp., Syrphus spp. and Episyrphus balteatus which are often much more abundant than their supposed models. These four taxa may still benefit from mimicry, but further research is needed to confirm this. With the possible exceptions of Eristalis pertinax and E. tenax , there is no strong evidence from north-west England in support of Waldbauer's hypothesis that mimics are rare when fledgling birds are abundant but may be commoner in autumn and spring. The data also indicate that mimics which closely resemble their models (specific mimics) are usually rarer than their models, whereas mimics with a less precise resemblance (non-specific mimics) are often commoner than models.     

The relationship between aphid infestations and oviposition by aphidophagous Syrphidae (Diptera)

Oviposition by aphidophagous Syrphidae varies with the size of aphid infestations and different syrphid species have different optimum aphid population sizes for oviposition. In one experiment using potted brussels sprout plants infested with Brevicoryne brassicae L., Platycheirus manicatus (Meig.) preferred about 100 aphids per plant, P. scutatus (Meig.) about 1000 aphids per plant and Syrphus ribesii (L.) about 2000 aphids per plant. These preferences were less clear in certain Platycheirus species (e.g. P. peltatus (Meig.)), than in Syrphus species (e.g. S. luniger Meig.). Once a plant has been selected for oviposition there may still be selection of suitable colonies on that plant. For a given aphid population, S. luniger preferred a small number of large aggregates to a larger number of smaller ones, whereas S. balteatus (Deg.) preferred the opposite. The tendency of the different species to select aphid populations of different sizes and distributions is likely to decrease interspecific competition.     

A study on pollen morphology of the genus Vitis L.

The pollen grains of 32 species, 1 subspecies and 4 varieties of Vitis and 1 species of Muscadinia were scanned with Hitachi S-800 scanning electron microscope and described in the present paper. In Vitis, the pollen morphology is found to be valuable for the identification of species, though it is of little significance in supraspecific classification and systematic consideration. The ornamentation of Vitis could be divided into four types: ( 1 ) The pollen grains are oblate or rarely spheroidal (P/E =0.97 ～ 1.08) in shape, with colpi wide enough to show dense granulae inside. The pollen grains of V. coignetiae Pull. Ex Planch. belong to this type. (2) The pollen grains are prolate to subprolate (P/E = 1.25 ～ 1.76 ), with sparse granulae within the colpi. This type includes l0 species, 1 subspecies and 3 varieties, such as V. hui Cheng etc. (3) The pollen grains of V. romanetii Roman du Caill. ex Planch. and V. davidii (Roman du Caill. ) Foex. var. ferruginea Merr. et Chun are prolate (P/E= 1.87 ～ 2.1), with colpi narrow and linear-shaped, but rhombic in the equational region. (4) The pollen grains are prolate or perprolate (P/E= 1.30 ～ 2. 0), with narrow and linear-shaped colpi. The species with this type of pollen grains are Muscadiniarotundifolia (Michx.) Small and 20 species of Vitis.