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.     

Zoogeographical analysis of the hoverfly fauna (Diptera, Syrphidae) of the northwestern Caucasus

The hoverfly fauna of the northwestern Caucasus comprises 257 species of 72 genera. They possess 32 types of ranges that could be united into 11 groups of ranges. Widespread species from the basis of the hoverfly fauna in the region: species with narrower ranges, those distributed in more than one zoogeographical region, and those distributed in more than one dominion, constitute 12.9%, 63.1%, and 24% of the fauna, respectively. The hoverfly fauna of the northwestern Caucasus is composed of nemoral species common for Europe and Mediterranean basin, and also by species widespread in the Palaearctic (92.9%). Autochthomous species constitute 5.5%, and species of the eastern Palaearctic origin, 1.6%. Hoverfly species with zoophagous larvae are the most widespread. The fraction of such species is low among species with narrowly local ranges (12.5%) and high among species with ranges including more than one dominion. Phytophagous species are wider represented among hoverflies with narrowly local ranges (44.7%) than among flies with interdominion ranges (8%). Ranges of dendrobiontic saprophagous species are determined by the presence of arboreal vegetation and no widespread polytopic hoverflies are found in this group. The distribution of mycetophagous species, commensals, and terrestrial saprophagous species is determined by the presence of fungi, ranges of some hymenopterans, and the presence of decaying plant remnants.     

The conservation requirements of an endangered hoverfly, Hammerschmidtia ferruginea (Diptera, Syrphidae) in the British Isles

The saprophagous larva of the endangered hoverfly, Hammerschmidtia ferruginea (Fallén, 1817) (Diptera, Syrphidae) develops in wet, decaying sap under the bark of dead trees and branches of aspen, Populus tremula L. (Saliaceae). However this breeding site is transient: wet decay builds up patchily over 1.5–2 years of the tree or branch dying and lasts for a further 1–3 years before the bark falls off. Between 1990 and 2006, H. ferruginea swung through a cycle of abundance when the number of localities where it was detected dropped from 13 to 5 and back to 8. Fluctuations in amounts of dead wood caused by winds and storms probably explain this population swing. When there are few dead trees and branches, H. ferruginea breeds in sap flows on live P. tremula trees and populations are maintained but at low levels. To prevent local extinctions during periods when fallen wood is scarce, breeding habitat can be supplemented by felling trees and branches. Fallen wood with wet decaying sap is also important as an assembly site for mate selection. Adult food plants include flowers of bird cherry Prunus padus, rowan Sorbus aucuparia and hawthorn Crategus monogyna. Adults were recorded dispersing up to 1 km but are probably able to move further than this.     

Seasonal dynamics of hoverfly (Diptera,Syrphidae) activity in the taiga zone of the Komi Republic

194 species of 56 genera of hoverflies are recorded from the taiga zone of the Komi Republic. The phenology of the hoverfly flying activity and temporal structure of their trophic and ecological assemblages have been investigated. Ten phenological groups are distinguished and characteristics of adult diets of each of these groups are given.     

Seasonal changes in a cactus‐hoverfly (Diptera: Syrphidae) network

1. We analysed a cacti‐syrphid community focusing on the exploitation of decaying cacti resources by the flies, i.e. if exploitation exhibited a nested topology as a structural pattern, or whether it was temporally random. If availability of cactus resources was predictable during the rainy season, it would then be exploited by a more structured community, or as resource availability would be unpredictable during the dry season, we should expect it to be exploited by a random community. 2. We registered 12 Copestylum species (9 in dry and 11 in rainy season). Four cactus species were recorded per season, three were shared between seasons. 3. The community of Copestylum larvae in the rainy season was not randomly assembled but highly nested, revealing a highly structured pattern of resource use. It exhibited a random organization for the dry season. The high nestedness value obtained for the rainy season suggests that factors along with competition must play a major role in determining community structure. 4. Succession in the cacti‐syrphid community mediated by microorganisms involved in necrosis is an important factor structuring nested subsets. The studied networks were small, which may limit the power of the analysis, and strong conclusions could also be limited.     

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.     

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.     

Effects of climate change on the distribution of hoverfly species (Diptera: Syrphidae) in Southeast Europe

Climate change presents a serious threat to global biodiversity. Loss of pollinators in particular has major implications, with extirpation of these species potentially leading to severe losses in agriculture and, thus, economic losses. In this study, we forecast the effects of climate change on the distribution of hoverflies in Southeast Europe using species distribution modelling and climate change scenarios for two time-periods. For 2041–2060, 19 analysed species were predicted to increase their areas of occupancy, with the other 25 losing some of their ranges. For 2061–2080, 55% of species were predicted to increase their area of occupancy, while 45% were predicted to experience range decline. In general, range size changes for most species were below 20%, indicating a relatively high resilience of hoverflies to climate change when only environmental variables are considered. Additionally, range-restricted species are not predicted to lose more area proportionally to widespread species. Based on our results, two distributional trends can be established: the predicted gain of species in alpine regions, and future loss of species from lowland areas. Considering that the loss of pollinators from present lowland agricultural areas is predicted and that habitat degradation presents a threat to possible range expansion of hoverflies in the future, developing conservation management strategy for the preservation of these species is crucial. This study represents an important step towards the assessment of the effects of climate changes on hoverflies and can be a valuable asset in creating future conservation plan, thus helping in mitigating potential consequences.     

Effect of hunger on yellow water trap catches of hoverfly (Diptera: Syrphidae) adults

1 An experiment was conducted in a winter wheat field using yellow water traps at crop height and at ground level, near to and distant from flowers, to test the hypothesis that such traps are seen as a source of food by flower‐feeding adult hoverflies and are therefore likely to selectively trap hungry individuals. Hoverflies caught in each trap were counted and identified and the amount of pollen in their guts was assessed. Ratios of numbers of hoverflies seen in the wheat crop to numbers caught in nearby traps were compared for the different treatments. 2 Most hoverflies were caught in crop‐high traps but they included a high proportion of individuals with empty guts. The taxa were: Episyrphus balteatus (De Geer), Metasyrphus corollae (F.) (these species together accounted for over 90% of the individuals trapped), with Sphaerophoria spp., Syrphus spp., Scaeva pyrastri (L.), Melanostomini and unidentified others making up the rest. 3 Significantly fewer hoverflies were captured in low than in high traps. The ratios of numbers trapped to numbers observed, in flower and in no flower treatments would be expected to be the same if the traps were not selective. This was supported for low traps. With high traps, however, there was a highly significant difference between the ratios (71.34 and 126 : 8, respectively). 4 Flies captured in high traps had less pollen in their gut than those captured in low ones. At each distance, more E. balteatus captured in high traps were in pollen category 1 (< 20 grains) than in any other category. The opposite state was seen in low trap catches, where most flies were in category 5 (> 5000 grains). Median pollen categories were 2 (21–200 grains) and 4 (1501–5000 grains) for flies caught in high and low traps, respectively. 5 The ecological selectivity of traps according to their height and the physiological condition of the targeted individuals is a problem likely to affect many trapping systems apart from the one described in this paper.     

Use of selected flowering plants in greenhouses to enhance aphidophagous hoverfly populations (Diptera: Syrphidae)

The addition of floral resources in a crop is the most commonly used conservation biological control strategy. The influence of additional floral resources on the abundance of aphidophagous syrphids has been studied in Mediterranean sweet-pepper greenhouses, in southeast Spain. Sweet alyssum and coriander were the plant species used as flowering plants, distributed in the greenhouse in several monospecific patches. In our first experiment the influence on syrphid pre-imaginal stages (larvae and pupae) was studied and adult stages were studied in a second experiment. A higher number of pre-imaginal syrphids was recorded in two replicated greenhouses where flowers were introduced, compared with two control greenhouses (without additional floral resources). To evaluate the effect on adults, 4 greenhouses were divided into 2 plots in each greenhouse and flowers were introduced in one plot per greenhouse. More hoverfly adults were observed in the plots where flowers had been introduced than in the control. The three most abundant syrphid species found were Eupeodes corollae, Episyrphus balteatus and Sphaerophoria rueppellii. Specimens from these species were dissected, and their pollen content was analysed to assess the food potential of the introduced flowers. The three syrphid species fed on pollen from both the flowering plants, as well as on sweet-pepper pollen. This conservation biological control strategy is an effective method to enhance native syrphid populations in Mediterranean sweet-pepper greenhouses.     

Food utilization of aphidophagous hoverfly larvae (Diptera: Syrphidae, Chamaemyiidae) on herbaceous plants in an urban habitat

We examined food utilization in a community of aphidophagous hoverfly larvae (Diptera: Syrphidae and Chamaemyiidae) in open lands in an urban habitat in central Japan for 3 years. The community consisted of 17 hoverfly species feeding on 20 aphid species occurring on 14 species of dominant herbaceous plants. In terms of larval prey preference, the dominant eight species of hoverfly were categorized into three groups: a polyphagous ‘generalist’ group consisting of four species,Episyrphus balteatus, Betasyrphus serarius, Syrphus vitripennis andSphaerophoria sp.; an oligophagous ‘specialist’ group consisting of three species,Metasyrphus hakiensis, Dideoides latus andParagus hemorrhous; andLeucopis puncticornis, which showed a preference for two aphid species on the plantTorilis scabra. The prey aphids of the second group have behavioral or morphological defense mechanisms that are effective for preventing attacks by generalist hoverflies; two prey aphids are aggressive toward generalist predators and the others are protected by ant-attendance. The specialist hoverflies seem to be adapted to overcome these defense mechanisms. The prey ranges overlapped little between the generalist and the specialist groups, while those within the generalist group overlapped greatly.     

Geographical patterns of hoverfly (Diptera,Syrphidae) functional groups in Europe: inconsistency in environmental correlates and latitudinal trends

Abstract 1. Relationships between species richness in higher taxa and large‐scale environmental variables have been widely studied over the past 15 years. Much less is known about how different functional groups (FGs) of species with similar biological and life‐history traits contribute to the overall trends, or how they differ in species‐richness patterns. 2. Multivariate analysis clustered 641 species of Syrphidae into eight FGs on the basis of 10 life‐history features, revealing feeding strategy as the main factor separating the groups. 3. Geographical trends in species richness and determinants of species richness within the FGs were compared across Europe. 4. Total species richness showed no latitudinal trend. However, the richness of individual FGs revealed variable relationships with latitude, including positive, negative, and hump‐shaped ones. This appeared to be related to how different environmental factors affected species richness within FGs. 5. Functional groups differed in their responses to the environmental variables. Annual temperature, evapotranspiration, and elevation span were the most important variables separating the FGs in ordination analysis. The multiple regression models showed further differences between FGs and their responses to the environment. 6. The FG approach revealed important inconsistencies in latitudinal diversity gradients and diversity‐climate relationships.     

长白山北坡不同海拔及干扰程度下访花食蚜蝇群落的差异

本研究采用垂直面取样法,对中国长白山北坡不同海拔及干扰程度下访花食蚜蝇群落的物种组成、多度、丰富度及多样性进行了调查研究。我们选择了不同垂直带的两个样地,分别位于800~1 100 m和1 750 ~2 150 m;每个样地选取3种生境类型,每一生境类型采样面积为20 m×20 m。252组调查数据表明,共采到访花食蚜蝇42种2 540个体。不同海拔生境访花食蚜蝇的组成、多度、丰富度及多样性的差异分析结果表明: 低海拔样地内不同生境访花食蚜蝇的群落相似性高于高海拔样地;低海拔样地访花食蚜蝇的多度高于高海拔样地,而物种丰富度低于高海拔样地;不同生境类型访花食蚜蝇多样性指数存在差异,但高海拔样地与低海拔样地多样性差异不显著;过渡林生境（岳桦林带）与次生林生境（针阔混交林带Ⅱ）中访花食蚜蝇的多样性较高。保护珍稀植物物种的人工植物花园访花食蚜蝇多样性仅次于过渡林生境（岳桦林带）与次生林生境（针阔混交林带Ⅱ）,保护作用显著。     

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.     

Saproxylic hoverflies benefit by modern forest management (Diptera: Syrphidae)

Saproxylic organisms are regarded as threatened in many European countries. Nevertheless, there have been some signals indicating a recent increase of saproxylic hoverflies in the Netherlands. This paper examines the change in occurrence of saproxylic hoverflies compared to other ecological groups of the same family. The trend analysis is based on the database of an extensive recording programme carried out in the Netherlands. The results show an increase in occupied grid cells for a majority of saproxylic species in relation to other ecological groups. This positive trend occurs among all five ecological subcategories of saproxylic hoverflies recognized in this paper. It is suggested that this increase could be attributed to the changes in the Dutch forests during the past 50 years. Important aspects of these changes are: the increase in forest area (including an increase in area of forest with large and very large trees), the increase in age of forests and the changes in forest management in favour of a more natural approach towards dead and ill trees. In the discussion the possible influence of climate change and the method of trend calculation are discussed.     

Mark recapture estimates of dispersal ability and observations on the territorial behaviour of the rare hoverfly,Hammerschmidtia ferruginea (Diptera,Syrphidae)

In order to effectively manage habitat for fragmented populations, we need to know details of resource utilisation, and the capacity of species to colonise unoccupied habitat patches. Dispersal is vital in maintaining viable populations in increasingly fragmented environments by allowing re-colonisation of areas in which populations have gone extinct. In the UK, the endangered aspen hoverfly Hammerschmidtia ferruginea (Fallén 1817) (Diptera, Syrphidae) depends on a limited and transient breeding habitat: decaying aspen wood Populus tremula L. (Salicaceae). Conservation management for H. ferruginea involves encouraging aspen expansion across Scotland, and ensuring retention, maintenance and continuity of dead wood where H. ferruginea has been recorded and in areas that may link populations. In order to do this effectively we need to know how far H. ferruginea can disperse. By taking advantage of the tendency of adults to group on decaying aspen logs, we estimated dispersal ability through mark recapture techniques. In the first year, 1,066 flies were marked as they emerged from aspen logs and 78 were re-sighted at artificially-placed decaying aspen logs up to 4 km from the release site. In the second year, of 1,157 individually marked flies, 112 were re-sighted and one was observed 5 km from the release site. Territorial behaviour was recorded at all (19) decaying aspen log locations. In total, 72 males were recorded defending territories, which overlapped with 68 % of recorded female oviposition sites. Among males only, wing length was positively associated with dispersal. While these results show H. ferruginea is capable of locating decaying logs up to 5 km away, most dispersing individuals (68 %) were recorded at 1 km, which should be taken into account in developing management protocols. If enough dead wood is available it should be distributed within a radius of 1–2 km, and where possible, as stepping-stones linking up aspen woodlands. We discuss the implications of our findings for the natural history of this species, and make recommendations for its conservation management.     

Restriction fragment length polymorphisms of different DNA regions as genetic markers in the hoverfly Episyrphus balteatus (Diptera: Syrphidae)

A polymerase chain reaction-restriction fragment length polymorphism (PCR-RFLP) analysis using mitochondrial (A+T-rich region; mtDNA) and genomic (zen-region; nDNA) DNA was performed on 182 female individuals of Episyrphus balteatus (DeGeer), a widespread aphidophagous hoverfly with supposed migratory behaviour. Specimens originated from 13 sampling sites in six European countries. The analyses revealed 12 and 18 haplotypes, respectively, for the two DNA types, several of them with a wide distribution, although seven and eight haplotypes, respectively, occurred only in one location. In contrast to other studies on mobile insects, the genetic diversity was relatively high. However, lack of population subdivision, low genetic distances between populations, the very high gene flow rates, and the complete lack of isolation by distance suggest that E. balteatus populations are largely connected and that there is an absence of large-scale geographic structuring. These results support the hypothesis that E. balteatus is a migratory hoverfly species, capable of moving over large distances. These findings related to the seasonal migrations of this species are discussed.     

黑带食蚜蝇Episyrphus balteatus De Geer的复眼结构及其调光机制

【目的】观察研究黑带食蚜蝇Episyrphus balteatus De Geer成虫复眼形态、小眼结构和不同光暗条件对小眼结构的影响,以明确其光视觉的结构基础和调光机制。【方法】利用组织切片法和扫描电镜等技术。【结果】1.复眼位于头部两侧,正面观呈半球形,占据除额颜外大部分头部。雄虫与雌虫单个复眼分别有约7 180个、7 230个小眼。各小眼面呈整齐排列的规则六边形。2.小眼由角膜及伪晶锥组成的屈光器、不同水平面分布的8个小网膜细胞及其特化形成的离散型视杆、屏蔽色素细胞和基膜等组成。小眼自远端至近端由主色素细胞和12个附属色素细胞围绕。3.随光暗条件的改变小眼内的附属色素细胞色素和基细胞细胞核沿小眼纵轴移动。光适应时,附属色素细胞色素颗粒沿小眼纵轴均匀分布,基细胞细胞核位于基膜上方。暗适应时,附属色素细胞色素颗粒向伪晶锥近端压缩,基细胞细胞核亦向远端移动,到达视杆中段。【结论】黑带食蚜蝇复眼精密的小眼排列形式和内部结构均显示了其强大的生理功能;屏蔽色素颗粒的移动是其复眼适应外界光环境变化的重要机制。本试验为进一步探究黑带食蚜蝇视觉结构和光调节机制,以及与其飞行行为间的关系提供了一定的理论基础。     

Internal labelling of the eggs of the aphidophagous hoverfly Episyrphus balteatus (De Geer) (Diptera: Syrphidae) with a fluorescent dye

A method was developed to label the egg load of Episyrphus balteatus (De Geer) (Diptera: Syrphidae) by feeding adult females with the fluorescent dye Rhodamine B. In a preliminary experiment aimed at selecting an appropriate dye concentration, gravid females reared in insectary conditions were provided for two successive days with a 50% honey-water solution enriched with Rhodamine B at concentrations of 0.1%, 0.01% and 0.001%. Eggs deposited on aphid-infested broad bean plants by syrphid females fed with Rhodamine B 0.01% showed the most intense marking. The higher dye concentration induced irregular marking among the eggs, presumably due to its repellent taste and consequently lower intake by the flies. A second series of trials focused on the persistence of the marker, prior and after oviposition, at the selected (optimal) dilution. A reduction in the marking degree of eggs was observed when the first egg deposition was delayed for one day after removing the dye from the cage. When marked females were allowed to deposit further batches of eggs the subsequent days, fewer eggs were laid and their fluorescence properties was reduced. Exposure to direct sunlight during 10 hours had no detectable effect on the fluorescence level of marked eggs deposited on a bean plant. The egg-marking technique examined in this study could be useful to determine the crop area accommodating eggs deposited by marked syrphid females migrating from a release point into cultivated fields.     

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.