Phylogenomics and biogeography of sawflies and woodwasps (hymenoptera, symphyta).

Phylogenomic approaches have recently helped elucidate various insect relationships, but large-scale comprehensive analyses on relationships within sawflies and woodwasps are still lacking. Here, we infer the relationships and long-term biogeographic history of these hymenopteran groups using a large dataset of 354 UCE loci collected from 385 species that represent all major lineages. Early Hymenoptera started diversifying during the Early Triassic ∼249 Ma and spread all over the ancient supercontinent Pangaea. We recovered Xyeloidea as a monophyletic sister group to other Hymenoptera and Pamphilioidea as sister to Unicalcarida. Within the diverse family Tenthredinidae, our taxonomically and geographically expanded taxon sampling highlights the non-monophyly of several traditionally defined subfamilies. In addition, the recent removal of Athalia and related genera from the Tenthredinidae into the separate family Athaliidae is supported. The deep historical biogeography of the group is characterised by independent dispersals and re-colonisations between the northern (Laurasia) and southern (Gondwana) palaeocontinents. The breakup of these landmasses led to ancient vicariance in several Gondwanan lineages, while interchange across the Northern Hemisphere has continued until the Recent. The little-studied African sawfly fauna is likewise a diverse mixture of groups with varying routes of colonization. Our results reveal interesting parallels in the evolution and biogeography of early hymenopterans and other ancient insect groups.

EUKARYOME: the rRNA gene reference database for identification of all eukaryotes.

Molecular identification of micro- and macroorganisms based on nuclear markers has revolutionized our understanding of their taxonomy, phylogeny and ecology. Today, research on the diversity of eukaryotes in global ecosystems heavily relies on nuclear ribosomal RNA (rRNA) markers. Here, we present the research community-curated reference database EUKARYOME for nuclear ribosomal 18S rRNA, internal transcribed spacer (ITS) and 28S rRNA markers for all eukaryotes, including metazoans (animals), protists, fungi and plants. It is particularly useful for the identification of arbuscular mycorrhizal fungi as it bridges the four commonly used molecular markers-ITS1, ITS2, 18S V4-V5 and 28S D1-D2 subregions. The key benefits of this database over other annotated reference sequence databases are that it is not restricted to certain taxonomic groups and it includes all rRNA markers. EUKARYOME also offers a number of reference long-read sequences that are derived from (meta)genomic and (meta)barcoding-a unique feature that can be used for taxonomic identification and chimera control of third-generation, long-read, high-throughput sequencing data. Taxonomic assignments of rRNA genes in the database are verified based on phylogenetic approaches. The reference datasets are available in multiple formats from the project homepage, http://www.eukaryome.org.

Palaearctic willow-catkin sawflies: a revision of the amentorum species group of Euura (Hymenoptera, Tenthredinidae).

The Euura amentorum species group is Holarctic, and in Europe it is most species-rich in the North. Their larvae develop entirely within the female catkins of Salix species: some species bore in the central stalk, whereas others live outside this and feed mainly on the developing seeds. Eight Palaearctic species are treated here as valid, and a key to these is provided. Males of five species are known. Two new species are described from northern Europe: Euura pohjola sp. n. and E. ursaminor sp. n. First records of E. itelmena (Malaise, 1931) from the West Palaearctic are presented. We propose seven new synonymies: Pontopristia montana Lindqvist, 1961 (junior secondary homonym in Euura) with Euura freyja (Liston, Taeger & Blank, 2009); Pontopristia brevilabris Malaise, 1921, Amauronematus fennicus Lindqvist, 1944, Pontopristia boreoalpina Lindqvist, 1961, Pontopristia punctulata Lindqvist, 1961, and Amauronematus pyrenaeus Lacourt, 1995 with Euura microphyes (Frster, 1854); and Pteronidea holmgreni Lindqvist, 1968 with Nematus umbratus Thomson, 1871. Lectotypes are designated for: Amauronematus fennicus Lindqvist, 1944, Nematus amentorum Frster, 1854, Nematus suavis Ruthe, 1859, Pontopristia brevilabris Malaise, 1921, Pontopristia itelmena Malaise, 1931, Pontopristia kamtchatica Malaise, 1931, Pontopristia lapponica Malaise, 1921, Pontopristia latiserra Malaise, 1921, Pontopristia romani Malaise, 1921, and Pristiphora amentorum var. nigripleuris Enslin, 1916. Many new host plant associations are recorded.

Cross-contamination and strong mitonuclear discordance in Empria sawflies (Hymenoptera, Tenthredinidae) in the light of phylogenomic data.

In several sawfly taxa strong mitonuclear discordance has been observed, with nuclear genes supporting species assignments based on morphology, whereas the barcode region of the mitochondrial COI gene suggests different relationships. As previous studies were based on only a few nuclear genes, the causes and the degree of mitonuclear discordance remain ambiguous. Here, we obtained genomic-scale ddRAD data together with Sanger sequences of mitochondrial COI and two to three nuclear protein coding genes to investigate species limits and mitonuclear discordance in two closely related species groups of the sawfly genus Empria. As found previously based on nuclear ITS and mitochondrial COI sequences, species are in most cases supported as monophyletic based on new nuclear data reported here, but not based on mitochondrial COI. This mitonuclear discordance can be explained by occasional mitochondrial introgression with little or no nuclear gene flow, a pattern that might be common in haplodiploid taxa with slowly evolving mitochondrial genomes. Some species in the E. immersa group are not recovered as monophyletic according to either mitochondrial or nuclear data, but this could partly be because of unresolved taxonomy. Preliminary analyses of ddRAD data did not recover monophyly of E. japonica within the E. longicornis group (three Sanger sequenced nuclear genes strongly supported monophyly), but closer examination of the data and additional Sanger sequencing suggested that both specimens were substantially (possibly 10-20% of recovered loci) cross-contaminated. A reason could be specimen identification tag jumps during sequencing library preparation that in previous studies have been shown to affect up to 2.5% of the sequenced reads. We provide an R script to examine patterns of identical loci among the specimens and estimate that the cross-contamination rate is not unusually high for our ddRAD dataset as a whole (based on counting of identical sequences in the immersa and longicornis groups, which are well separated from each other and probably do not hybridise). The high rate of cross-contamination for both E. japonica specimens might be explained by the small number of recovered loci (~1000) compared to most other specimens (>10 000 in some cases) because of poor sequencing results. We caution against drawing unexpected biological conclusions when closely related specimens are pooled before sequencing and tagged only at one end of the molecule or at both ends using a unique combination of limited number of tags (less than the number of specimens).

A review of West Palaearctic Hoplocampa species, focussing on Sweden (Hymenoptera, Tenthredinidae).

Fourteen Hoplocampa species have been recorded in the West Palaearctic. We provide an illustrated key to these species, together with H. tadshikistanica, which is so far only known from Tadshikistan, but could occur in the West Palaearctic. The suitability of genetic sequencing for identification, particularly of larvae, is discussed. COI barcoding reliably distinguishes all European species which have been sampled (only H. phantoma lacks data), except for H. fulvicornis and H. minuta, which can be identified using nuclear sequences. Distributions in the Fennoscandian countries are outlined, with particular reference to Sweden. Hoplocampa chrysorrhoea is recorded for the first time in Scandinavia, from southern Sweden. Lectotypes are designated for twelve nominal taxa: Allantus ferrugineus Panzer, 1802, Hoplocampa chrysorrhoea var. nigrita Enslin, 1914, H. fabricii W. F. Kirby, 1882, H. oertzeni Konow, 1888, H. pectoralis Thomson, 1871, Hylotoma ferruginea Fabricius, 1804, Tenthredo alpina Zetterstedt, 1838, T. brevis Klug, 1816, T. chrysorrhoea Klug, 1816, T. crataegi Klug, 1816, T. plagiata Klug, 1816, and T. rutilicornis Klug, 1816. Hoplocampa minuta forma dudai Gregor, in Gregor Bata, 1942 is a new synonym of H. fulvicornis (Panzer, 1801).

The West Palaearctic Pseudodineura and Endophytus species (Hymenoptera, Tenthredinidae).

Six valid species of Pseudodineura are now recognised as occurring in the West Palaearctic, and the only described species of the related genus Endophytus. Larvae of all species are leaf-miners in Ranunculaceae. An identification key to adults is provided, followed by species commentaries which include summarised data on taxonomy, larval host plants, and distribution, with particular reference to Sweden. Whereas identification of some specimens using morphological characters may not be possible, each species apparently has a distinct COI barcode sequence. Pseudodineura heringi (Enslin, 1921) is a new junior synonym of P. parvula (Klug, 1816). Pseudodineura mocsaryi Zombori, 1976 and P. scaligera Zombori, 1979 are new junior synonyms of P. clematidisrectae Hering, 1935. Lectotypes are designated for: Dolerus minutus Hartig, 1837, Pelmatopus clematidis Hering, 1932, P. enslini Hering, 1923, P. heringi Enslin, 1921, and P. mentiens var. konowi Enslin, 1921.

The West Palaearctic Dineura species, focussing on Sweden (Hymenoptera, Tenthredinidae).

Four Dineura species are now considered to occur in the West Palaearctic, including northern Europe, but D. parcivalvis has not been found in Scandinavia. Dineura pullior Schmidt Walter, 1995 is treated as a new junior subjective synonym of D. virididorsata (Retzius, 1783). An illustrated key to adults is presented. Lectotypes are designated for seven nominal taxa: Dineura stilata var. virilis Enslin, 1918, Dineura testaceipes var. nigriventris Enslin, 1915, Dineura virididorsata var. dorsalis Enslin, 1915, Nematus posticus Förster, 1854, Nematus xanthocerus Hartig, 1840, Nematus xanthopus Zaddach, 1876, and Tenthredo (Allantus) stilata Klug, 1816. Distributions in the Fennoscandian countries are outlined, with particular reference to Sweden.

The West Palaearctic genera of Nematinae (Hymenoptera, Tenthredinidae).

Keys to adults and larvae of the genera of West Palaearctic nematine sawflies are presented. Species of some of the smaller genera are keyed, and their taxonomy, distribution, and host plants reviewed, with a geographic focus on north-western Europe, particularly Sweden. Dinematus Lacourt, 2006 is a new junior subjective synonym of Pristiphora Latreille, 1810, resulting in the new combination Pristiphora krausi (Lacourt, 2006) for the type species of Dinematus. Hemichroa monticola Ermolenko, 1960 is a new junior subjective synonym of Hemichroa australis (Serville, 1823). Lectotypes are designated for Tenthredo opaca Fabricius, 1775, Mesoneura opaca var. nigerrima Enslin, 1914, Mesoneura opaca var. obscuriventris Enslin, 1914, Nematus hypogastricus Hartig, 1837, Nematus alnivorus Hartig, 1840, Leptopus rufipes Förster, 1854, Nematus protensus Förster, 1854, and Platycampus luridiventris var. pleuritica Enslin, 1915. A phylogenetic analysis based on four genes (mitochondrial COI and nuclear NaK, POL2, and TPI) supports the current generic classification.

Identification of sawflies and horntails (Hymenoptera, 'Symphyta') through DNA barcodes: successes and caveats.

The 'Symphyta' is a paraphyletic assemblage at the base of the order Hymenoptera, comprising 14 families and about 8750 species. All have phytophagous larvae, except for the Orussidae, which are parasitoids. This study presents and evaluates the results of DNA barcoding of approximately 5360 specimens of 'Symphyta', mainly adults, and 4362 sequences covering 1037 species were deemed of suitable quality for inclusion in the analysis. All extant families are represented, except for the Anaxyelidae. The majority of species and specimens are from Europe, but approximately 38% of the species and 13% of the specimens are of non-European origin. The utility of barcoding for species identification and taxonomy of 'Symphyta' is discussed on the basis of examples from each of the included families. A significant level of cryptic species diversity was apparent in many groups. Other attractive applications include the identification of immature stages without the need to rear them, community analyses based on metabarcoding of bulk samples and association of the sexes of adults.

Host plants of Empria sawflies (Hymenoptera, Tenthredinidae) in Japan include Rhododendron (Ericaceae).

New host plant records are given for six Empria species from Japan. They are Rosa multiflora [Rosaceae] for E. honshuana Prous & Heidemaa, 2011, Rubus sp. [Rosaceae] for E. japonica Heidemaa & Prous, 2011, Geum japonicum and G. calthifolium var. nipponicum [Rosaceae] for E. loktini Ermolenko, 1971, Rosa multiflora, Potentilla indica and probably Rubus parvifolius [Rosaceae] for E. quadrimaculata Takeuchi, 1952, Rhododendron molle subsp. japonicum [Ericaceae] for E. takeuchii Prous & Heidemaa, 2011, and Geum japonicum and Filipendula camtschatica [Rosaceae] for E. tridentis Lee & Ryu, 1996. This is the first record of Ericaceae as a host plant of Empria. The mode of host shifts in the evolution of Empria is inferred by using a phylogenetic hypothesis proposed by Prous et al. (2011a).

The larva and prepupa of Eupareophora exarmata (Thomson, 1871) (Hymenoptera, Tenthredinidae).

BACKGROUND: Of the two known Eupareophora species, more is known about the larva  and bionomics of the Nearctic E. parca, than the rarely recorded West Palaearctic E. exarmata. NEW INFORMATION: The last instar larva and prepupa of E. exarmata is illustrated and briefly described. In Germany its host is Fraxinus excelsior.

Sawfly taxa (Hymenoptera, Symphyta) described by Edward Newman and Charles Healy.

Type specimens of seven nominal species of sawfly described by Edward Newman and one by Charles Healy were studied. This material is housed in the Oxford University Museum of Natural History, United Kingdom. The following new synonymies are proposed (valid names in parentheses): Hartigia Schiødte, 1839 (Phylloecus Newman, 1838), Cephus helleri Taschenberg, 1871 (Phylloecus faunus Newman, 1838) and Euura gallae Newman, 1837 (Euura mucronata (Hartig, 1837)). The type species of Euura Newman, 1837 and Euura subgenus Gemmura E. L. Smith, 1968 belong to the same taxonomic species, Euura mucronata (Hartig, 1837), so that these genus group names become new synonyms. Lectotypes are designated for Phyllotoma tormentillae Healy, 1868, Fenusa ianthe Newman, 1837, Fenusa parviceps Newman, 1837, Selandria pallida Newman, 1837 and Phylloecus faunus Newman, 1838. 26 new combinations are proposed for species formerly placed in Hartigia and here transferred to Phylloecus, and 4 original combinations are re-instated as valid.

Review of the sawfly genus Empria (Hymenoptera, Tenthredinidae) in Japan.

The following eleven Empria species are reported from Japan: Empria candidata (Fallén, 1808), Empria japonica Heidemaa & Prous, 2011, Empria liturata (Gmelin, 1790), Empria loktini Ermolenko, 1971, Empria plana (Jakowlew, 1891), Empria quadrimaculata Takeuchi, 1952, Empria rubicola Ermolenko, 1971, Empria tridens (Konow, 1896), Empria tridentis Lee & Ryu, 1996, Empria honshuana Prous & Heidemaa, sp. n., and Empria takeuchii Prous & Heidemaa, sp. n. The lectotypes of Poecilosoma pallipes Matsumura, 1912, Empria itelmena Malaise, 1931, Tenthredo candidata Fallén, 1808, and Tenthredo (Poecilostoma) hybrida Erichson, 1851 are designated. Empria itelmena Malaise, 1931, syn. n. is synonymized with Empria plana (Jakowlew, 1891). Poecilosoma pallipes Matsumura, 1912, previously assigned to Empria, is transferred to Monsoma, creating Monsoma pallipes (Matsumura, 1912), comb. n. Results of phylogenetic analyses using mitochondrial (COI) and nuclear (ITS1 and ITS2) sequences are also provided.