Gunilla Ståhls

First phylogeny of predatory flower flies (Diptera, Syrphidae, Syrphinae) using mitochondrial COI and nuclear 28S rRNA genes: conflict and congruence with the current tribal classification

The family Syrphidae (Diptera) is traditionally divided into three subfamilies. The aim of this study was to address the monophyly of the tribes within the subfamily Syrphinae (virtually all with predaceous habits), as well as the phylogenetic placement of particular genera using molecular characters. Sequence data from the mitochondrial protein‐coding gene cytochrome c oxidase subunit I (COI) and the nuclear 28S ribosomal RNA gene of 98 Syrphinae taxa were analyzed using optimization alignment to explore phylogenetic relationships among included taxa. Volucella pellucens was used as outgroup, and representatives of the tribe Pipizini (Eristalinae), with similar larval feeding mode, were also included. Congruence of our results with current tribal classification of Syrphinae is discussed. Our results include the tribe Toxomerini resolved as monophyletic but placed in a clade with genera Ocyptamus and Eosalpingogaster. Some genera traditionally placed into Syrphini were resolved outside of this tribe, as the sister groups to other tribes or genera. The tribe Bacchini was resolved into several different clades. We recovered Paragini as a monophyletic group, and sister group of the genus Allobaccha. The present results highlight the need of a reclassification of Syrphinae.

COI barcodes for identification of Merodon hoverflies (Diptera, Syrphidae) of Lesvos Island, Greece

DNA barcoding has become a useful system for linking different biological life stages, and for identification of species within a known taxonomic framework. In this study, we generated mitochondrial DNA COI barcodes using adult specimens of all 22 species of the hoverfly genus Merodon (Diptera, Syrphidae) occurring on Lesvos island (Greece). The generated COI barcodes could well discriminate between all Merodon taxa of Lesvos, except for M. loewi and M. papillus that shared the same haplotype, despite their clear morphological differences. In addition, the barcodes revealed two cases of hitherto unknown morphologically cryptic species close to M. avidus and M. nigritarsis, respectively. Because only few successful rearings of immature stages of Merodon hoverflies are available, the larval host plant remains unknown for these phytophagous taxa. The obtained COI barcode library for the Merodon spp. of Lesvos will constitute a tool to link any unknown immature stages with already known species, and thus provide important life‐history information and promise for ecological studies.

Molecular phylogeny of Allograpta (Diptera, Syrphidae) reveals diversity of lineages and non-monophyly of phytophagous taxa

Phylogenetic relationships of genera Allograpta, Sphaerophoria and Exallandra (Diptera, Syrphidae) were analyzed based on sequence data from the mitochondrial protein-coding gene cytochrome c oxidase subunit I (COI) and the nuclear 28S and 18S ribosomal RNA genes. The three genera are members of the subfamily Syrphinae, where nearly all members feed as larvae on soft-bodied Hemiptera and other arthropods. Phytophagous species have recently been discovered in two subgenera of Allograpta, sg Fazia and a new subgenus from Costa Rica. Phylogenetic analyses of the combined datasets were performed using parsimony, under static alignment and direct optimization, maximum likelihood and Bayesian inference. Congruent topologies obtained from all the analyses indicate paraphyly of the genus Allograpta with respect to Sphaerophoria and Exallandra. Exallandra appears embedded in the genus Sphaerophoria, and both genera are placed within Allograpta. The distribution of phytophagous taxa in Allograpta indicates that plant feeding evolved at least twice in this group.

Two new species of the genus Merodon Meigen 1803 (Diptera: Syrphidae) from the island of Lesvos (Greece), in the eastern Mediterranean

Abstract Descriptions are given of two new species of Merodon Meigen (Diptera: Syrphidae) from the island of Lesvos (Greece): Merodon papilius Vujić, Radenković & Pérez-Bañon n. sp. from the ruficornis group species and Merodon sapphous Vujić, Pérez-Bañon & Radenković n. sp. from the aureus group. In addition to classical morphological characters, partial sequences of the mitochondrial COI gene are generated for each taxon as DNA ‘barcodes’. Main characteristics of the species habitats are reported. The zoogeographical significance of these endemic taxa to the biodiversity of the island is discussed.

Generic revision and species classification of the Microdontinae (Diptera, Syrphidae)

Abstract With 552 species group names available (excluding misspellings), the Microdontinae constitute the smallest of the three subfamilies of Syrphidae. Paradoxically, this subfamily is taxonomically the least organized of the three: 388 species names were previously classified in a single genus, Microdon Meigen, 1803. The present paper introduces a new generic classification of the Microdontinae, relying partly on the results of phylogenetic analyses of morphological and molecular data as published in other papers, and partly on examination of primary type specimens of 347 taxa, plus additional material, and original descriptions. A total number of 67 genus group names (excluding misspellings) are evaluated, redescribed, diagnosed and discussed, with several implications for their taxonomic status. Of these, 43 names are considered as valid genera, 7 as subgenera, 17 as synonyms. Two generic names (Ceratoconcha Simroth, 1907, Nothomicrodon Wheeler, 1924) are left unplaced, because they are known from immature stages only and cannot be reliably associated with taxa known from adults. The following 10 new genera are described by Reemer: Domodon, Heliodon, Laetodon, Menidon, Mermerizon, Metadon, Peradon, Piruwa, Sulcodon and Thompsodon. A key to all genera, subgenera and species groups is given. A total number of 26 new species are described in the following genera: Archimicrodon Hull, 1945, Ceratrichomyia Séguy, 1951, Domodon, Furcantenna Cheng, 2008, Heliodon, Indascia Keiser, 1958, Kryptopyga Hull, 1944, Masarygus Brèthes. 1908, Mermerizon, Metadon, Microdon, Paramixogaster Brunetti, 1923, Piruwa, Pseudomicrodon Hull, 1937, Rhopalosyrphus Giglio-Tos, 1891, and Thompsodon. New lectotypes are designated for Ceratrichomyia behara Séguy, 1951 and Microdon iheringi Bezzi, 1910. A total number of 267 new combinations of species and genera are proposed. New synonyms are proposed for 19 species group names. Three replacement names are introduced for primary and secondary junior homonyms: Microdon shirakii nom. n. (= Microdon tuberculatus Shiraki, 1968, primary homonym of Microdon tuberculatus de Meijere, 1913), Paramixogaster brunettii nom. n. (= Mixogaster vespiformis Brunetti, 1913, secondary homonym of Microdon vespiformis de Meijere, 1908), Paramixogaster sacki nom. n. (= Myxogaster variegata Sack, 1922, secondary homonym of Ceratophya variegata Walker, 1852). An attempt is made to classify all available species names into (sub)genera and species groups. The resulting classification comprises 454 valid species and 98 synonyms (excluding misspellings), of which 17 valid names and three synonyms are left unplaced. The paper concludes with a discussion on diagnostic characters of Microdontinae.

Systematics and taxonomy of the ruficornis group of genus Merodon Meigen (Diptera: Syrphidae)

We revise the ruficornis group of species of Merodon Meigen (Diptera, Syrphidae) providing an illustrated key, a discussion of taxonomic characters and a morphological diagnosis. A total of 18 species from the ruficornis group are treated including distributional data. Descriptions are provided for seven new species: M. gallicus Vujić & Radenkovićsp.n., M. hoplitis Hurkmans sp.n., M. lamellatus Vujić & Radenkovićsp.n., M. nigripodus Vujić & Hayat sp.n., M. ovaloides Vujić & Radenkovićsp.n., M. ponticus Vujić & Radenkovićsp.n., and M. turcicus Vujić & Hayat sp.n. The taxon M. auripes Sack, is redefined and a neotype designated. Lectotypes are designated for M. graecus Loew; M. planiceps Loew and M. crymensis Paramonov. The monophyly and systematic position of this species group was assessed based on parsimony and maximum likelihood analyses of mitochondrial COI and nuclear 28S rDNA sequences. A very high level of endemism was observed in the ruficornis species group. Among 18 taxa, 12 are limited‐range endemics, present in few mountain areas or in a small part of the total range of the group. These endemics clearly show the importance of geographic isolation in the process of speciation.

Investigating plant—pollinator relationships in the Aegean: the approaches of the project POL-AEGIS (The pollinators of the Aegean archipelago: diversity and threats)

Summary Worldwide, there is a well-documented crisis for bees and other pollinators which represent a fundamental biotic capital for wild life conservation, ecosystem function, and crop production. Among all pollinators of the world, bees (Hymenoptera: Apoidea) constitute the major group in species number and importance, followed by hover flies (Diptera: Syrphidae). The Aegean constitutes one of the worlds hotspots for wild bee and other pollinator diversity including flies (mainly hover flies and bee flies), beetles, and butterflies. Despite this advantage, our present knowledge on Greek pollinators is poor, due to a lack of focused and systematic research, absence of relevant taxonomic keys, and a general lack of taxonomic experts in the country. As a result, assessments of pollinator loss cannot be carried out and the causes for the potential pollinator loss in the country remain unknown. Consequently, the desperately needed National Red Data list for pollinators cannot be compiled. This new research (2012–2015) aims to contribute to the knowledge of the pollinator diversity in Greece, the threats pollinators face, as well as the impacts these threats may have on pollination services. The research is conducted in the Aegean archipelago on > 20 islands and several mainland sites in Greece and Turkey. Prime goals are: i. the assessment of bee and hover fly diversity (species, genetic); ii. their pollination services; and iii. the effects of climate change, grazing, intensive bee-keeping, fires, electromagnetic radiation on bee diversity and ecology, as well as on plant-pollinator networks. At the same time, this research contributes to the taxonomic capital in Greece and the Eastern Mediterranean, focusing on the creation of the first identification keys for pollinators, the training of new scientists, as well as the enrichment and further development of the Melissotheque of the Aegean, a permanent reference collection of insect pollinators established at the University of the Aegean.

Is the mega-diverse genus Ocyptamus (Diptera, Syrphidae) monophyletic? Evidence from molecular characters including the secondary structure of 28S rRNA

Phylogenetic relationships between two New World Syrphinae taxa (Diptera, Syrphidae), i.e. the highly diverse genus Ocyptamus and the large genus Toxomerus, were analysed based on molecular characters. The monophyly of both taxa was tested and the taxonomic status of included subgenera and species groups was examined. Toxomerus constitutes the monogeneric tribe Toxomerini with more than 140 described species, while Ocyptamus (tribe Syrphini) is a very diverse genus (over 300 spp.) with multiple recognised subgenera and species groups. Sequence data from three gene regions were used: the mitochondrial protein-coding gene cytochrome c oxidase subunit I (COI) and the nuclear 28S and 18S ribosomal RNA genes. The secondary structure of two expansion segments (D2, D3) of the ribosomal 28S RNA gene is presented for the family Syrphidae and used for the first time in a multiple sequence alignment. Molecular data were analysed using parsimony, maximum likelihood and Bayesian inference. Toxomerus was always recovered as monophyletic within Ocyptamus, and relationships to other New World taxa such as Salpingogaster (Eosalpingogaster) were well-supported. Only the subgenera and species groups of Ocyptamus were consistently recovered as monophyletic lineages, thus the apparent non-monophyly of Ocyptamus demands reclassification of this clade.

Cheilosia vernalis (Diptera, Syrphidae) Complex: Molecular and Morphological Variability

Genus Cheilosia (Diptera, Syrphidae) with more than 300 Palaearctic species is the most speciose group of hoverflies in Europe. One of the most widespread taxa with unclear taxonomic status is Cheilosia vernalis (Fallen, 1817). This species shows great morphological variation in external morphological characters. A subdivision of the taxon has been suggested, but the male terminalia of the different morphological forms appear identical. The aim of this study was to integrate the information from mitochondrial COI sequence data with re-evaluated morphological characters for exploring species boundaries and for revealing phylogenetic relationships of C. vernalis and closely related species. We sequenced a 698 bp fragment of the 3′-end of the mitochondrial gene cytochrome oxidase subunit I of 43 specimens agreeing with traditional morphological interpretation of C. vernalis (Diptera, Syrphidae), sampled from a broad geographic range in Europe and single representatives of 10 species of the vernalis—melanura complex. Parsimony analysis of the molecular data showed C. vernalis specimens grouped in three different lineages. The C. vernalis lineage presents 8 different mitochondrial haplotypes. We found no congruence between these haplotypes and diagnosable morphological characteristics of the adult fly specimens supporting species subdivision. Based on re-examination of type material two new synonyms are proposed: C. rotundiventris Becker, 1894 and C. ruficollis Becker, 1894 are junior synonyms of C. vernalis (Fallen, 1817). New diagnostic morphological character for C. reniformis Hellen, 1930 is added and first distribution data of this species in Europe are given. Taxonomic status of C. longifila Becker, 1894 remains unresolved.

Phylogenetic relationships and taxonomic ranking of pipizine flower flies (Diptera: Syrphidae) with implications for the evolution of aphidophagy

The taxonomic rank and phylogenetic relationships of the pipizine flower flies (Diptera: Syrphidae: Pipizini) were estimated based on DNA sequence data from three gene regions (COI, 28S and 18S) and 111 adult morphological characters. Pipizini has been treated as a member of the subfamily Eristalinae based on diagnostic adult morphological characteristics, while the larval feeding mode and morphology is shared with members of the subfamily Syrphinae. We analysed each dataset, both separately and combined, in a total evidence approach under maximum parsimony and maximum likelihood. To evaluate the influence of different alignment strategies of rDNA 28S and 18S genes on the resulting topologies, we compared the topologies inferred from a multiple alignment using fast Fourier transform (MAFFT) program with those topologies resulting from aligning the secondary structure of these rDNA genes. Total evidence analyses resolved pipizines as a sister group of the subfamily Syrphinae. Although the structural alignment and the MAFFT alignment differed in the inferred relationships of some clades and taxa, there was congruence in the placement of pipizines. The homogeneous morphology of the Pipizini clade in combination with their unique combination of characters among the Syrphidae suggest a change of rank to subfamily. Thus, we propose to divide Syrphidae into four subfamilies, including the subfamily Pipizinae stat. rev.