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.

     

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.     

Biology of Bactrocera carambolae (Diptera: Tephritidae) on four hosts

Bactrocera carambolae is a quarantine pest found in Brazil, restricted to the states of Amapá, Pará and Roraima. This fruit fly can potentially cause extensive socioeconomic and environmental damage in the country, if it disperse into areas where fruit is grown for exporting. The objective of this work was to study the biology of B. carambolae on fruits of Averrhoa carambola L. (Oxalidaceae), Psidium guajava L. (Myrtaceae), Spondias mombin L. (Anacardiaceae) and Eugenia stipitata McVaugh (Myrtaceae). The following parameters were investigated: duration of egg-larva, pupal, egg-adult, pre-oviposition, oviposition and post-oviposition periods, pupal weight and viability, sex ratio, fecundity, fertility and longevity. All parameters except pupal weight, oviposition and post-oviposition period, egg fertility and sex ratio were influenced by the host plant on which the larvae were reared. The carambola fruit fly completes its development on all those hosts studied here, with the highest fecundities on A. carambola and P. guajava.     

Diversity increases with elevation: empidine dance flies (Diptera,Empididae) challenge a predominant pattern

Tropical mountain forests are important reservoirs of biodiversity. They are usually species‐rich and often support endemic species making them prime targets for conservation effort. The aim of this study was to investigate elevational patterns of species diversity and phenology to provide a meaningful understanding of insects’ spatio‐temporal distributions along a tropical gradient of elevation. Our study focuses on the Empidinae communities (Diptera, Empididae) from Doi Inthanon (north Thailand, gradient 400–2556 m asl), sampled during two entire years (2006–2007, 2014). This group of insects is more diverse in temperate localities than in the tropics and we found that: (1) increase in altitude and latitude has a similar effect, so that the diversity of our model increases with elevation; (2) the phenology is strongly influenced by seasonality with a peak of diversity occurring during the transition between the dry and rainy seasons; (3) there is no phenological shift in the diversity peak with elevations; and (4) the species composition changes along the altitudinal gradient and through the year (high beta diversity). The increase in diversity with increasing elevation and the peak of diversity occurring at the transitional period both strongly coincide with abiotic factors, the decreasing temperature and the arrival of the monsoon, respectively.     

Factors Influencing Bug Diversity (Insecta: Heteroptera) in Semi-Natural Habitats

We investigated the abundance and species richness of heteropteran bugs and explored environmental factors which influence bug diversity in three types of semi-natural habitats (wildflower areas, extensively used meadows, extensively grazed pastures). To cover this topic, it is essential to know how much the relatively young wildflower areas contribute to biodiversity compared with well-established extensive meadows and pastures. Total bug species richness and phytophagous bug species richness were significantly higher in wildflower areas and meadows than in pastures. In wildflower areas, we found the highest number of zoophagous bug species and species overwintering in the egg-stage. Species overwintering as adults were most abundant in meadows. Total number of bug species as well as species richness in either trophic groups and overwintering strategies were significantly positively correlated with vegetation structure. Except for overwintering strategies, the same was true to bug abundance. The bug community based on the number of individuals per species was significantly explained by flower abundance and vegetation structure, accounting for 18.4 and 16.8% of the variance, respectively. Our results indicate that vegetation structure and flower abundance are key factors for bug species richness, abundance and bug species composition. Since wildflower areas and meadows clearly increased bug species richness and contained several specialised bug species that did not occur in pastures, we recommend the promotion of wildflower areas and extensively used meadows in order to restore both high heteropteran diversity and overall insect biodiversity in agricultural landscapes.     

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.     

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.     

First use of pheromones to detect phenology patterns and density relationships of pine sawflies in German forests

1 Monitoring studies of pine sawflies with pheromone traps were performed for the first time in Germany. Pheromone traps baited with species‐specific pheromone substances were installed in pine forests at different locations in Bavaria, Brandenburg and Lower Saxony during two years. 2 It was possible to track the flight phenology of Diprion pini, Gilpinia pallida and Neodiprion sertifer reliably and to get information about the number of generations of these species in 1997 and 1998. 3 A clear relationship between trap catch and population density could not be detected, but qualitative changes in trap catch caused by different density levels were observed. 4 For D. pini, trap catches were different among endemic populations of different forest types. Furthermore, catches of males reflected the results from the regular cocoon collections by foresters during the previous winter. 5 For N. sertifer, trap catches in endemic populations were well separated from trap catches on sites with higher sawfly densities. However, no significant correlation between trap catch and sawfly density or defoliation level could be found. These results suggest that the efficacy of the pheromone traps probably varied with biological features (sex ratio, density level, immigration) of the particular population.     

Visits to Manipulated Flowers by Episyrphus balteatus (Diptera: Syrphidae): Partitioning the Signals of Petals and Anthers

In a number of previous studies attention has been directed to the selection on corolla dimensions by pollinator preference, but anthers may also be a signal. This experiment examined the relative importance of petals and anthers in the attraction of male Episyrphus balteatus (Diptera, Syrphidae) to individual oil-seed rape flowers (Bracus rapae oleifera). Rape flowers have four petals and six anthers. Nine treatments were created, of flowers with four, two, or zero petals and six, three, or zero anthers. Twenty males were tested for preference among the experimental flowers by recording the sequence and duration of their visits (which were all for feeding) to the flowers. The number of first visits and the duration of visits on each flower were significantly related to the number of anthers but not the number of petals. There was no correlation between petal area and pollen load of control flowers. It is suggested that Syrphidae may not always attend to petals, and variation in preference of different pollinator taxa should be taken into account when discussing the evolution of floral characters.     

Mimicry in hoverflies (Diptera: Syrphidae): a field test of the competitive mimicry hypothesis

Although most studies on the evolution of mimicry and warningcoloration in insects have considered predators as the majorselective force, it is possible that competition for food resourcescould also facilitate selection for these conspicuous signals.For example, when warningly colored social wasps visit flowers,then they frequently behave aggressively toward heterospecifics,and they also attack and feed on other flying insects. Underthese conditions, a resemblance to a wasp might provide a mimetichoverfly with improved access to floral resources by reducingthe frequency with which it is disturbed by other pollinators.We experimentally evaluated whether wasp-like colors and patternswere important in preventing other flower visitors from sharingthe same flower resource, using pairwise presentations of bothnatural and artificial prey in the field. Flower visitors weremore likely to visit unoccupied flowers compared with the flowerspinned with either natural or artificial specimens in 2 plantspecies with different inflorescences. However, flower visitorsdid not show a significantly reduced rate of visitation to flowerspinned with specimens bearing wasp-like colors and patternscompared with the flowers occupied by similar-sized specimensthat were nonmimetic. Overall, we found no compelling evidencein this study to support the contention that wasp-like warningsignals of hoverflies prevent other flower visitors from sharingflower resources, although insects showed a greater tendencyto avoid visiting flowers pinned with a wasp compared with flowerspinned with a nonmimetic fly.     

Distribution and abundance of aphidophagous hoverflies (Diptera: Syrphidae) in wildflower patches and field margin habitats

1 The spatial and temporal variations in aphidophagous syrphid abundance were recorded over two seasons in wildflower resource patches sown in a winter barley crop and associated field margins. Standard census techniques and sticky board trapping were used to assess numbers of syrphids, whilst weekly flower head counts were used to quantify the floral resources available in each of the patches. 2 The field margin supported a greater diversity and density of syrphids than the within‐crop wildflower patches, despite having a relatively lower flower head density. Presumably this was in response to other resources that field margins offer, namely additional aphid resources, shelter from predation, lekking sites and suitable flight‐paths. 3 The commonest species of syrphid, Episyrphus balteatus, demonstrated a very positive habitat association with the field margin and was rarely reported in the field patches. Therefore, it may be an unsuitable candidate for the biological control of aphids via augmentation of numbers using non‐host resources. 4 Patch size and shape had little effect on the spatial distributions of syrphids, probably because of the adult syrphids' high mobility. 5 Of greater influence was the number of flowers contained in each habitat patch. Typically, patches with higher numbers of flowers had significantly greater aggregations of hoverflies. Habitat manipulation by the provision of flowers in patches seems to increase the local density of hoverflies. Further work is necessary to establish the importance of flower density in enhancing the control of pest populations.     

Determination of phylogenetic position ofPipizini (Diptera: Syrphidae): based on molecular biological and morphological data

Based on the sequence analysis of 5.8S subunit and internal transcribed spacers (ITS) of ribosomal RNA gene (rDNA), the molecular phylogenetic tree of representative species of Pipizini and three groups of Syrphidae with different feeding habits (seven species belong to six genera) was constructed. Meanwhile, the phylogenetic tree of tribes (including Pipizini and other 17 tribes of Syrphidae) was constructed using morphological characteristics of adults and larvae and the number of chromosomes. Both the results show that the relationship between Pipizini and predatory groups is closer than that between Pipizini and saprophagous groups. So it is suggested that Pipizini be transferred from Milesiinae to Syrphinae.     

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.     

A new black fly species of the subgenus Gomphostilbia (Simulium: Simuliidae: Diptera) from India

Simulium (Gomphostilbia) agasthyamalaiense sp. nov. is described based on adults, pupae and mature larvae from a medium-flowing stream of Southern Western Ghats, India. This new species is placed in the Simulium batoense species-group of the subgenus Gomphostilbia Enderlein. This new species is characterized in the female by a scutum with three brownish-black longitudinal vittae and the hind basitarsus 5.7 times as long as wide; in the female by the large facets of upper eye with 20 vertical columns and 19 horizontal rows; in the pupa the respiratory gill with medium-long common basal stalk; and in the larva arrowhead-shaped postgenal cleft. Taxonomic notes are provided for this new species and it can be distinguished from closely related species of S. (G.) peteri. Keys are constructed to distinguish this species from ten species of the batoense species-group recorded in India.www.zoobank.org/urn:lsid:zoobank.org:pub:1D1CFEE5-A762-4F68-9EAE-9E31133146C0     

长白山北坡阔叶红松林及其次生白桦林高等植物物种多样性比较

应用Ｓｉｍｐｓｏｎ和Ｓｈａｎｎｏｎ多样性指数及其相应的均匀度,对阔叶红松林及其次生白桦林的高等植物物种多样性进行了对比研究．结果表明,对于木本植物,次生白桦林物种多样性高于阔叶红松林;而对于草本植物,情况正好相反．但在次生白桦林中,无论木本植物还是草本植物,占优势的物种都是一些常见种,而许多阔叶红松林中的珍稀或特有物种在次生白桦林中数量很少,有些甚至完全消失．对群落各种群多度分布的检验表明,２种森林类型中,无论木本植物还是草本植物,各种群的多度分布都遵从对数级数分布．     

The role of odour and visual cues in the pan-trap catching of hoverflies (Diptera: Syrphidae)

Coloured pan traps are frequently used to attract and catch insects, such as in the monitoring of populations of beneficial insects in classical or conservation biological control. They are also used in the evaluation of the recovery of insect populations after disturbance and in many other situations where an estimate of relative insect numbers is required. However, the fact that traps may be visible to the insects over a considerable distance can influence the interpretation of catch data. This difficulty may arise, for example, if traps along a transect can attract insects from some or all of the other transect positions. This study compared the effect of different coloured traps on attraction and catch of hoverflies. The hypothesis was that completely yellow traps would attract hoverflies from a distance, while traps that were green outside and yellow inside would catch fewer flies because only those from above or near the trap can see the yellow stimulus. A subsidiary hypothesis was that rose water would enhance hoverfly capture rates. For the two main hoverfly species captured, Melanostoma fasciatum (Macquart) and Melangyna novaezelandiae (Macquart), significantly more individuals were caught in completely yellow traps than in yellow and green or in completely green traps. Moreover, the addition of rose water increased the number of hoverflies caught significantly. It is suggested that if a measure of hoverfly numbers relating to a particular distance along a transect is required, consideration should be given to the ability of hoverflies to see yellow traps from a distance. The use of traps that are green outside would more accurately reflect the local abundance of hoverflies. If higher trap catches of hoverflies are needed for statistical purposes, rose water can enhance catches.     

Determination of phylogenetic position of Pipizini (Diptera: Syrphidae): based on molecular biological and morphological data

Based on the sequence analysis of 5.8S subunit and internal transcribed spacers (ITS ) of ribosomal RNA gene (rDNA), the molecular phylogenetic tree of representative species of Pipizini and three groups of Syrphidae with different feeding habits (seven species belong to six genera) was constructed. Meanwhile, the phylogenetic tree of tribes (including Pipizini and other 17 tribes of Syrphidae) was constructed using morphological characteristics of adults and larvae and the number of chromosomes. Both the results show that the relationship between Pipizini and predatory groups is closer than that between Pipizini and saprophagous groups. So it is suggested that Pipizini be transferred from Milesiinae to Syrphinae.     

Abundance and host-seeking activity of the biting midge,Forcipomyia taiwana (Diptera: Ceratopogonidae)

The abundance and host-seeking activity of the biting midge, Forcipomyia taiwana (Shiraki), were measured in Central Taiwan at 418 locations with low, medium, and high densities using a human-bait method. Abundance estimates at 10 min after sampling commenced were comparable to longer sampling periods of 20 min, allowing a shorter 10 min sampling period to be used throughout. Host-seeking F. taiwana females were only active during daylight and biting activity did not occur after sunset. The diurnal activity of host-seeking females changed seasonally, beginning at around 07:00–08:00 during the spring equinox, summer solstice, and autumnal equinox, whereas activity began an hour later in winter. Most females were collected during 10:00–16:00, with a peak abundance during 13:00–15:00. Blood-sucking activity of female F. taiwana were positively correlated with ambient temperature with a minimum threshold of 18 °C. When different years, seasons, trapping times, environmental factors, and female adults trapped were analyzed using multiple regression model, the results showed that all these factors contributed to the regression model and the determination coefficient (R²) of this multiple regression model was 0.81.