Optical releasers of the innate proboscis extension in the hoverfly Eristalis tenax L. (Syrphidae,Diptera)

Freshly emerged, inexperienced imagos of the hoverfly Eristalis tenax L. extend their proboscis towards small, yellow colour stimuli, such as anther parts and artificial floral guides. The releasing of this behaviour, which is adapted to pollen feeding, was investigated in behavioural tests using white, UV-reflecting artificial flowers with four small screens illuminated with test stimuli serving as artificial floral guides. The releasing of the innate proboscis extension was tested using monochromatic test lights. Within an intensity range from approx. 5·10¹¹ to approx. 10¹⁴ quanta · cm^-2· s^-1, the flies extended their proboscis only towards green and yellow test lights (approx. 520–600 nm). The inhibition of the innate proboscis extension was tested using mixed light stimuli composed of a yellow monochromatic reference light (560 nm, 10¹³ quanta·cm^{-2 -1}) and of a monochromatic test light. When the reference light was mixed with ultraviolet or blue test lights, the releasing of the innate proboscis extension was strongly inhibited, whereas admixing green/yellow light slightly promoted it; admixing red light had no effect. The results indicate that the releasing of the innate proboscis extension is mediated by the photoreceptor type R8y. Other receptor types which could cause the inhibition of the proboscis reaction are discussed.     

Intra-puparial development in the hoverflies Eristalinus aeneus and Eristalis tenax (Diptera: Syrphidae)

The intra-puparial development of 150 pupae of Eristalinus aeneus (Scopoli, 1763) and Eristalis tenax (Linnaeus, 1758) was analyzed. Individuals were obtained from the sixth laboratory generation kept under artificial rearing conditions at the facilities of the University of Alicante (Spain). The experiment was conducted at 25 ± 1°C temperature, 50 ± 5% relative humidity, and 12:12 hr (L:D) of photoperiod. Groups of 10 pupae were collected every 6 hr over 48 hr, after that period, pupae were collected daily until the adult emergence. They were fixed in 5% formic acid and preserved in 70% ethanol. Fixed pupae were dissected and photographed. The chronology and morphological changes that take place during the intra-puparial development in both species were analyzed and compared. Five phases were observed: prepupa, before 6 hr; cryptocephalic pupa, between 6 and 24 hr; phanerocephalic pupa, between 24 and 30 hr; pharate adult, after 30 hr; and the adult imago, restricted to the very end of the development process just before adult emergence. In total, the intra-puparial development lasted 189 ± 4 hr in E. aeneus and 192 ± 3 hr in E. tenax, with the pharate adult the longest phase (some 81% of the total developmental time). These data can be used to develop accurate cold storage protocols during artificial rearing of both pollinator species, avoiding critical events during the development and increasing survival.     

Phenotypic variation in colour pattern and seasonal plasticity in Eristalis hoverflies (Diptera: Syrphidae)

Abstract.

• 1 An examination of phenotypic variation in colour pattern was carried out on four Eristalis hoverfly species using museum material.
• 2 The amount of phenotypic variation varied substantially among the species with E.arbustorum being the most variable. The other species showed a wide colour pattern range but less variation within that range (E.abusivus and E.nemorum), or a narrow range of colour variation (E.horticola).
• 3 Sexual colour dimorphism was apparent in all four species, but most pronounced in E.abusivus and E.nemorum.
• 4 There were good phenotype-season relationships shown by both sexes in all species, except for female E.abusivus and E.nemorum, with paler insects being more abundant during the warmer summer months.
• 5 Female, but not male, E.arbustorum collected at inland sites were on average paler than those collected at coastal sites. This observation is considered with respect to temperature during the developmental stages.
• 6 The function of colour plasticity in hoverflies is discussed with reference to the need to maintain optimal thermal conditions for activity.

     

Immature stages of some aphidophagous syrphid flies of India (Insecta, Diptera, Syrphidae)

Immature stages, viz. egg, larva and puparium, of six species of the economically most important syrphid flies, Scaeva latimaculata (Brunetti), Ischiodon scutellaris (Fabricius), Episyrphus altemans (Macquart), Sphaerophoria Indiana Bigot, Metasyrphus confrater (Wiedemann), and M. latilunulatus (Collin), have been studied. These species are voracious feeders on three aphid species, Lipaphis erysimi (Kaltenbach), Myzus persicae (Sulzer) and Brevicoryne brassicae (L.), which are serious pests of mustard, Brassica campestris L.     

Pollination of the orchidEpipactis thunbergii by syrphid flies (Diptera: Syrphidae)

The behavior of visitors to the flowers of the orchidEpipactis thunbergii was studied, with special attention to the role of the epichile in the pollination process. Only four species of syrphid flies legitimately pollinated the flower, among whichSphaerophoria macrogaster was regarded as the most effective pollinator. The movable epichile, possessing a furrow at its base, played a critical role in the pollination process: it threw the syrphid fly onto the stigmatic surface when both sides of the basal slanting surface of the furrow were presumably pressed in the direction of the hypochile by the fore (and middle) legs of the retreating syrphid fly. At this moment, the fly received a set of pollinia on the thorax.     

Various stages in the life cycle of syrphid flies (Eristalis tenax; diptera: Syrphidae) as potential mechanical vectors of pathogens causing mycobacterial infections in pig herds

We defined the role of the syrphid fly Eristalis tenax in the survival and transmission of mycobacteria in pigs. The conditionally pathogenic mycobacterial (CPM) species Mycobacterium chelonae was isolated from 10 % of liquid dung samples, and both M. chelonae and another CPM species M. fortuitum were isolated from 7 (78 %) of the examined E. tenax larvae collected from the same location. Mycobacteriosis of the lymph nodes of pigs from 3 infected farms was caused by M. avium subsp. avium, M. avium subsp. hominissuis, and M. fortuitum. M. avium subsp. avium and M. avium subsp. hominissuis of identical genotype and serotypes and M. fortuitum were isolated from 7 (1.9 %) larvae, 2 (7.4 %) puparia, and one (1.6 %) imago. The count of colony forming units isolated from larval skin covering (pouch) was higher (p < or = 0.01) than that isolated from the internal organs of larvae. These results showed the potential for E. tenax larvae to spread mycobacteria throughout pig herds and the surrounding environment.     

The selective use of floral resources by the hoverfly Episyrphus balteatus (Diptera: Syrphidae) on farmland

The use of flowers of non-crop plants by the syrphid Episyrphus balteatus in margins of cereal fields was assessed quantitatively. G-tests and the ‘C’ index of Murdoch (1969) were used to compare the use and abundance of twenty-seven plant species by adult flies from June to August in the UK. The flowers of several plant species were used selectively by hoverflies throughout the season while the relative importance of others changed with date. The prospects for managing the field-margin flora to enhance biocontrol by hoverflies is discussed.     

Behavioural mimicry of honeybees (Apis mellifera) by droneflies (Diptera: Syrphidae: Eristalis spp.)

Droneflies (Syrphidae: Eristalis spp. resemble honeybees (Apis mellifera) in appearance and have often been considered to be Batesian mimics. This study used a focal watch technique in order to compare the foraging behaviour of droneflies Eristalis tenax, Eristalis pertinax, Eristalis arbustorum and Eristalis nemorum) whilst they were feeding on patches of flowers with the behaviour of honeybees and other hymenopterans and dipterans. It was found that, on a range of plant species, the time droneflies spent on individual flowers and the time spent flying between them was more similar to that of honeybees than to the times of other hymenopterans and dipterans. These results suggest that dronefly behaviour has evolved to become more similar to that of honeybees and they support the hypothesis that droneflies are Batesian mimics.     

Demography and population parameters of two species of eristaline flower flies (Diptera,Syrphidae, Eristalini)

The importance of eristaline flies (Diptera, Syrphidae, Eristalini) as pollinators in natural ecosystems and for agricultural crops is well known. However, in-depth studies on the life cycle of most of these species have yet to be carried out. The aim of this research was to study the life cycle of Eristalis tenax (Linnaeus, 1758) and Eristalinus aeneus (Scopoli, 1763) in order to improve the current rearing system employed at the University of Alicante. The results were analysed using the age-stage, two-sex life table method. As one of the main results, the mean duration of the life history of E. tenax and E. aeneus was 46.06 and 65.12 days (d), respectively. The most critical step for both species was found at the beginning of the larval stage (first instar), when the highest mortality was recorded. Population parameters were also analysed and compared. The intrinsic rate of increase (r), the finite rate of increase (λ), the net reproductive rate (R₀) and the mean generation time (T) were 0.09 (d⁻¹), 1.09 (d⁻¹), 42.11 offspring and 40.97 d, in the case of E. aeneus and 0.05 (d⁻¹), 1.05 (d⁻¹), 23.13 offspring and 59.23 d, in the case of E. tenax. These results indicate that the current rearing system is more efficient for E. aeneus, which displays a faster population growth. However, some modifications need to be implemented to improve the production of E. tenax.     

Field boundaries as barriers to movement of hover flies (Diptera: Syrphidae) in cultivated land

Field boundaries play an important role as refuges, food sources and corridors for invertebrates and vertebrates, and increasing farmland fragmentation impacts on these functions. However, hedgerows and other structures can also impede dispersal by flying insects. The current work uses the pollen of Phacelia tanacetifolia in hover fly guts as a marker to assess hover fly movement in farm landscapes. In the United Kingdom and New Zealand, Phacelia pollen was found in the guts of Ephisyrphus balteatus and Metasyrphus corollae (United Kingdom) and Melanostoma fasciatum (New Zealand) at distances up to 200 m from the source, when there were no barriers between the flowers and the traps used to catch the flies. The rate of decline over distance in the proportion of flies containing pollen was similar for the two countries. The extent to which four replicated field boundary types impeded hover fly movement was tested using post-and-wire fences, lines of poplars (Populus spp.) with gaps, dense poplars and controls (no potential barriers). Phacelia was planted on one side of each boundary, and along the centre of the control plots. The relative presence of the pollen in flies on both sides of the barriers showed that both types of poplar boundary restricted the movement of the flies, but the fence had no effect. In a separate experiment, gravid females of M. fasciatum were captured at a greater height on a shade-cloth fence than were non-gravid females and males. The implications of this work include the functioning and persistence of metapopulations and the influence of field boundaries on population recovery of beneficial invertebrates following pesticide-induced mortality. If field boundaries contribute to a temporal asynchrony between pest and natural enemy populations, this needs to be considered along with the well-established roles of boundaries as refugia for, and sources of, beneficial arthropods.     

Comparative characteristics of the wing apparatus and flight of Syrphid flies (Diptera: Syrphidae)

Comparative analysis of the wing apparatus and flight in nine species of flower flies (Syrphidae) has been performed. Data on the flight velocity, aerodynamic force, wing-beat frequency, stroke amplitude and stroke plane angle, wing area, body mass and volume, as well as correlations between these parameters at the intraspecific and family levels, have been obtained. Based on the obtained data, the subfamilies Syrphinae and Eristalinae have been compared.     

Systematics and evolution of predatory flower flies (Diptera: Syrphidae) based on exon-capture sequencing

Flower flies (Diptera: Syrphidae) are one of the most species-rich dipteran families and provide important ecosystem services such as pollination, biological control of pests, recycling of organic matter and redistributions of essential nutrients. Flower fly adults generally feed on pollen and nectar, but their larval feeding habits are strikingly diverse. In the present study, high-throughput sequencing was used to capture and enrich phylogenetically and evolutionary informative exonic regions. With the help of the baitfisher software, we developed a new bait kit (SYRPHIDAE1.0) to target 1945 CDS regions belonging to 1312 orthologous genes. This new bait kit was successfully used to exon capture the targeted loci in 121 flower fly species across the different subfamilies of Syrphidae. We analysed different amino acid and nucleotide data sets (1302 loci and 154 loci) with maximum likelihood and multispecies coalescent models. Our analyses yielded highly supported similar topologies, although the degree of the SRH (global stationarity, reversibility and homogeneity) conditions varied greatly between amino acid and nucleotide data sets. The sisterhood of subfamilies Pipizinae and Syrphinae is supported in all our analyses, confirming a common origin of taxa feeding on soft-bodied arthropods. Based on our results, we define Syrphini stat.rev. to include the genera Toxomerus and Paragus. Our divergence estimate analyses with beast inferred the origin of the Syrphidae in the Lower Cretaceous (125.5–98.5 Ma) and the diversification of predatory flower flies around the K–Pg boundary (70.61–54.4 Ma), coinciding with the rise and diversification of their prey.     

Diurnal activity patterns in hoverflies (Diptera, Syrphidae)

Abstract. 1. Time-budgets are constructed from censuses of hoverflies. Larger species spend less time in flight.
2. Most species are active under similar conditions of light intensity, but a shade-tolerant (Melanostoma scalare Fabr.) and a sun-loving species (Metasyrphus corollae Fabr.) were identified.
3. Large species become active (i.e. move, by flying or feeding) at lower temperatures than small ones, except M.scalare, which is tolerant to cool temperatures. This means that, M.scalare apart, the order of species appearing during the day is size-dependent, largest first.
4. Previous claims about the diel periodicity of syrphids can be reconciled by considering the importance of individual thermal balance.
5. Tolerance to low temperatures in small hoverflies is associated with feeding on anemophilous pollen; this may indicate a proline-fuelled flight.     

The influence of the development rates of immature stages of Eristalis arbustorum (Diptera; Syrphidae) on adult abdominal colour pattern

Abstract. The abdominal colour pattern of Eristalis arbustorum (L.) shows seasonal variation, with animals emerging in spring being darker than those emerging during summer. One of the most important environmental cues influencing the abdominal colour pattern is developmental temperature. An experiment was carried out to establish which period during the immature stages was important in determining final adult colour pattern. The results indicate a good relationship between the length of the pupal period and both the extent of the paler markings on the abdomen and the grey score of these markings. The length of the larval period did not have any effect on adult colour pattern.     

Functional richness of local hoverfly communities (Diptera, Syrphidae) in response to land use across temperate Europe

Environmental change is not likely to act on biodiversity in a random manner, but rather according to species traits that affect assembly processes, thus, having potentially serious consequences on ecological functions. We investigated the effects of anthropogenic land use on functional richness of local hoverfly communities of 24 agricultural landscapes across temperate Europe. A multivariate ordination separated seven functional groups based on resource use, niche characteristics and response type. Intensive land use reduced functional richness, but each functional group responded in a unique way. Species richness of generalist groups was nearly unaffected. Local habitat quality mainly affected specialist groups, while land use affected intermediate groups of rather common species. We infer that high species richness within functional groups alone is no guarantee for maintaining functional richness. Thus, it is not species richness per se that improves insurance of functional diversity against environmental pressures but the degree of dissimilarity within each functional group.