ore JHR 40: I-69 (2014) JOURNAL OF | *0eerviewed opevaccets ural doi: 10.3897/JHR.40.7442 ME Hymenopter a http://jhr.pensoft.net/ The inerational Society of Hymenoptersts. RESEARCH The genera of Nematinae (Hymenoptera, Tenthredinidae) Marko Prous'’, Stephan M. Blank', Henri Goulet’, Erik Heibo*, Andrew Liston', Tobias Malm°, Tommi Nyman?®, Stefan Schmidt®, David R. Smith’, Hege Vardal?, Matti Viitasaari?, Veli Vikberg'®, Andreas Taeger' | Senckenberg German Entomological Intitute, Eberswalder Strafse 90, 15374 Miincheberg, Germany 2 Department of Zoology, Institute of Ecology and Earth Sciences, University of Tartu, Vanemuise 46, 51014 Tartu, Estonia 3 Agriculture and Agri-Food Canada, 960 Carling Ave, ON KIA 0C6 Ottawa, Ca- nada 4 Ento Consulting, Rypeveien 34 A, 3420 Lierskogen, Norway § Department of Biology, University of Eastern Finland, PO. Box 111, FI-80101 Joensuu, Finland 6 Hymenoptera Section, Bavarian State Collec- tion of Zoology, Miinchhausenstrafse 21, 81247 Miinchen, Germany 7 Systematic Entomology Laboratory, Agricultural Research Service, U. S. Department of Agriculture, c/o National Museum of Natural History, Smithsonian Institution, RO. Box 37012, MRC 168, Washington, DC 20013-7012 8 Department of Zoology, Swedish Museum of Natural History, Box 50007, SE-104 05 Stockholm, Sweden 9 Alkutie 41E, 00660 Helsinki, Finland \0 Liinalammintie 11 as. 6, FI-14200 Turenki, Finland Corresponding author: Marko Prous (mprous@senckenberg.de) Academic editor: Gavin Broad | Received 7 March 2014 | Accepted 29 April 2014 | Published 2 October 2014 http:/lzo0bank.org/D14B3522-C772-4792-8F3D-DC59E047EEDO Citation: Prous M, Blank SM, Goulet H, Heibo E, Liston A, Malm T, Nyman T, Schmidt $, Smith DR, Vardal H, Viitasaari M, Vikberg V, Taeger A (2014) The genera of Nematinae (Hymenoptera, Tenthredinidae). Journal of Hymenoptera Research 40: 1-69. doi: 10.3897/JHR.40.7442 Abstract Recent phylogenetic studies on Nematinae based on DNA sequences have shown extensive incongruencies with current nomenclature of genus-group taxa. Here, we expand previous DNA sequence datasets based on three genes (Col, Cytb, and EF-1«), to include a fourth (NaK) and more genera. The analyses largely confirm the previous findings, particularly the existence of two well-supported large clades, Euura and Pristiphora, together comprising more than 75% of the species of Nematinae. Basal relationships within these two clades remain poorly resolved, mirroring the difficulties in delimiting genera based on morphology. In addition, a moderately supported small clade, Nematus, is found. ‘The relationships between the Euura, Pristiphora, and Nematus clades are uncertain. Therefore, to stabilize the nomenclature we treat these clades as genera. This taxonomic treatment results in numerous new combinations of species names. The following synonymies are Copyright Marko Prous et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 2 Marko Prous et al. / Journal of Hymenoptera Research 40: 1-69 (2014) proposed for the available genus-group names. Synonyms of Euura Newman, 1837: Cryptocampus Hartig, 1837, Evura Agassiz, 1848, Pontania Costa, 1852, syn. n., Epitactus Forster, 1854, syn. n., Amauronematus Konow, 1890, syn. n., Holcocneme Konow, 1890, syn. n., Pachynematus Konow, 1890, syn. n., Holcocnema Schulz, 1906, syn. n., Holcocnemis Konow, 1907, syn. n., Pteronidea Rohwer, 1911, syn. n., Pontopristia Malaise, 1921, syn. n., Brachycoluma Strand, 1929, syn. n., Decanematus Malaise, 1931, syn. n., Pikonema Ross, 1937, syn. n., Phyllocolpa Benson, 1960, syn. n., Fitelius Kontuniemi, 1966, syn. n., Gemmura E.L. Smith, 1968, Eupontania Zinovjev, 1985, syn. n., Larinematus Zhelochovtsev, 1988, syn. n., Polynema- tus Zhelochovtsev, 1988, syn. n., Bacconematus Zhelochovtsev, 1988, syn. n., Alpinematus Lacourt, 1996, syn. n., Epicenematus Lacourt, 1998, syn. n., Kontuniemiana Lacourt, 1998, syn. n., Lindqvistia Lacourt, 1998, syn. n., Luea Wei and Nie, 1998, syn. n., and Tubpontania Vikberg, 2010, syn. n. Synonyms of Nematus Panzer, 1801: Craesus Leach, 1817, Hypolaepus W.F. Kirby, 1882, and Paranematus Zinovjev, 1978. Synonyms of Pristiphora Latreille, 1810: Diphadnus Hartig, 1837, Lygaeonematus Konow, 1890, Microne- matus Konow, 1890, Gymnonychus Marlatt, 1896, Neopareophora MacGillivray, 1908, syn. n., Neotomosteth- us MacGillivray, 1908, Dineuridea Rohwer, 1912, Sala Ross, 1937, Pristola Ross, 1945, syn. n., Nepionema Benson, 1960, syn. n., Melastola Wong, 1968, syn. n., Sharliphora Wong, 1969, Oligonematus Zhelochovt- sev, 1988, Lygaeotus Liston, 1993, Lygaeophora Liston, 1993, and Pristicampus Zinovjev, 1993, syn. n. Varna Ross, 1937, syn. n. is treated as a synonym of Dineura Dahlbom 1835. Stauronematus Benson, 1953 is treated as a separate genus from Pristiphora. Names of 20 species-group taxa are junior secondary homonyms when combined with Euura. Replacement names are proposed for these. To facilitate the identification of Nematinae genera, we provide an illustrated key to the 31 extant genera of world Nematinae. Keywords Sawflies, taxonomy, phylogeny, key, new synonyms, new combinations, replacement names Introduction The Nematinae is the second-largest subfamily within the Tenthredinidae (Taeger et al. 2010). With about 1250 species, it is surpassed only by the Tenthredininae (about 1700 species) (Taeger and Blank 2011). In the Electronic World Catalogue of Symphyta (ECat- Sym) (Taeger and Blank 2011), 48 genera of Nematinae are currently recognised, but there is still no wider consensus on how many genera should be recognised, and how these should be delimited. Also lacking are comprehensive keys for identification of the genera. Only some regional keys are available (e.g. Benson 1958; Zhelochovtsev and Zinovjev 1988; Goulet 1992) which have to be combined with numerous other publications (To- gashi 1964; Taeger 1989; Zinovjev 1993; Wei 1998a; b; Wei and Nie 1998; Zinovjev 2000; Lacourt 2006; Wei and Nie 2008) to cover most of the genera of the world. Many (sub) genera are based on few morphological characters. Furthermore, some of the char- acter states lack discrete differences, making recognition of genera (and therefore species) difficult. Recent phylogenetic analyses based on DNA sequence data (Nyman et al. 2006; 2010) have shown that many genera (especially within the so-called ‘higher Nematinae’) are not monophyletic, indicating a need for a taxonomic revision. Here we expand these previous phylogenetic analyses (based on at most three genes: Col, Cytb, and EF-1«) with the addition of one nuclear gene (NaK) and more genera to provide a new generic classifi- cation of Nematinae. The new classification seems to offer the best prospect of promoting The genera of Nematinae (Hymenoptera, Tenthredinidae) 3 future stability of nomenclature. An extensively illustrated key to genera is also provided. This can be considered as a starting point for the revision of roughly half of the ca. 600 West Palaearctic nematine species, a project funded by the Swedish Taxonomy Initiative (STI). Unfortunately, the key does not solve all the problems in identifying genera. Per- haps the biggest drawback is our current inability to unequivocally separate Euura from Nematus based on morphology. The forthcoming keys to species of individual genera should remove some such remaining ambiguities in identifying genera. Material and methods Phylogenetic analyses We extracted DNA from larval and adult samples stored in 99.5% ethanol at —20 °C by using the DNeasy Tissue Kit (Qiagen, Valencia, CA). Sequence data were collected from the mitochondrial genes Cytochrome oxidase I (Col; 810 bp) and Cytochrome b (Cytb; 718 bp), and the nuclear genes Elongation factor-l« (EF-1«; two exons of the F2 copy; 777 bp in all) and Sodium—potassium adenosine triphosphatase (NaK; 997 bp). PCR am- plification and sequencing of Col, Cytb, EF-1«, and NaK were done as described earlier (Nyman et al. 2000; Nyman et al. 2006; Leppanen et al. 2012). New sequences have been deposited in GenBank under accession numbers KJ434795—KJ434930. Col and Cytb sequences of Monocellicampa pruni were extracted from a partial mitochondrial genome available in GenBank (JX566509; Wei et al. 2013b). In some cases, unpublished barcode sequences (Col; 658 bp) from the BOLD database (http://www.boldsystems.org/) were also included in the analysis to maximize representation of type species of genera. We constructed two four-gene alignments. The first contains 134 specimens and 3302 base pairs with little missing data. Most specimens in this dataset have sequenc- es from all four genes, but 19 specimens are missing one gene and 7 specimens are missing two genes. In order to examine the relationships between type species of the genus-group names, a second dataset of 79 specimens and 3537 bp was constructed. This alignment is longer because of the addition of the 658-bp barcode region of Col (Hebert et al. 2003): 423 bp of this region overlaps with the 5’ end of the 810-bp Col portion used in the first dataset. When the overlapping region was identical between two different conspecific specimens, a single composite terminal taxon was created to minimize the missing cells in the dataset. In this second dataset, 21 specimens are miss- ing one gene, 8 specimens are missing two genes, and 15 specimens are missing three genes. Four non-type species, Pristicampus incisus (Lindqvist, 1970), Paranematus tu- lunensis (Vikberg, 1972), Craterocercus fraternalis (Norton, 1872), and Susana annulata D.R. Smith, 1969, were added to the dataset, because the amount of sequence data for the respective type species (Pristicampus arcticus (Lindqvist, 1959) [422 bp of the Col barcode region], Paranematus wahlbergi (Thomson, 1871) [658 bp of Col], Craterocer- cus obtusus (Klug, 1816) [658 bp of Col], and Susana cupressi Rohwer and Middleton, 1932 [997 bp of NaK]) was insufficient to reliably estimate their phylogenetic position. 4 Marko Prous et al. / Journal of Hymenoptera Research 40: I-69 (2014) When we exclude genus names which are based on fossils, or are taxonomically unplaced, there are 78 genera based on different type species. Of these, 62 type species have DNA data. Of the remaining 16 type species, most can be associated using mor- phology with species for which DNA data is available, but Anhoplocampa, Armenocam- pus, Dinematus, Katsujia, Megadineura, Nescianeura and Renonerva still lack DNA data. We performed Bayesian phylogenetic analyses in MrBayes v. 3.2.2 (Ronquist et al. 2012) and maximum likelihood (ML) analyses in RAxML v. 7.6.6 (Stamatakis 2006; Stamatakis et al. 2008) using the CIPRES Science Gateway (Miller et al. 2010) at http://www.phylo.org/index.php/portal/. The dataset was partitioned by genes, and the best-fitting DNA substitution model for each gene was selected using jModel- Test 2.1.4 (Darriba et al. 2012), which uses PhyML (Guindon and Gascuel 2003) for likelihood calculations. Model selection was done using five substitution schemes (including parameters for base frequencies, gamma-distributed rate variation across sites (G) with four categories, and a proportion of invariable sites (I), altogether 40 dif- ferent models) on the basis of the Akaike Information Criterion (AIC). The best-fitting model for all four genes was GIR+1+G, which was used in MrBayes. In the RAxML runs, proportion of invariable sites (I) was not used as recommended in the manual of the program. In MrBayes, we used default priors, and each of the four partitions was allowed to have its own unlinked substitution model. We ran two parallel runs having four incrementally heated chains for 20 million generations, while sampling trees from the current cold chain every 1000 generations. We discarded 5000 trees sampled prior to reaching chain stationarity as a burn-in from both runs, and the remaining 15001 trees were used to calculate a 50% majority consensus rule tree, showing all groupings with posterior probability more than 0.5. In RAxML a separate GI'R+G model was employed for each gene, and node supports were evaluated based on 500 bootstrap replicates. The root of the phylogenetic trees on Figs 2—5 was placed between Nemati- nae and other representatives of Tenthredinidae in the dataset. For the subfamilial phylogenetic relationships and placement of Nematinae within Tenthredinidae, see Malm and Nyman (2014). Preparation of the key and morphological terminology For the generic key, we studied most of the type species of the genus-group taxa, and ad- ditional species when necessary (excluding fossils). Only 5 out of 80 type species of the available genus-group names given for extant taxa were not available for study: Anhop- locampa fumosa Wei, 1998, Caulocampus necopinus Zhelochovtsev, 1941 (type species of Armenocampus Zinovjev, 2000), Messa hortulana Leach, 1817 (unplaced name), Pris- tiphora varipes Lepeletier, 1823 (Stevenia Brullé, 1846; unplaced name), and Renonerva fumosa Wei and Nie, 1998. Except for Messa hortulana and Pristiphora varipes, the pub- lished descriptions and figures were sufficient to include them in the key. Morphological terminology is based on Goulet (1992) and Viitasaari (2002) with small modifications to the terminology of wing veins and cells (Fig. 1). The term campaniform sensilla The genera of Nematinae (Hymenoptera, Tenthredinidae) 5 Figure |. Wing veins (left) and cells of Hoplocampa chrysorrhoea (Klug, 1816). Veins: 1A, 2A, 3A — first, second, and third anal vein; C — costa; Cu — cubitus; Cul — cubitus 1; Culb — cubitus 1b; cu-a - cubito- anal cross-vein; M — medius; m-cu — medio-cubital cross-vein; Im-cu, 2m-cu — first, second medio- cubital cross-vein; R — radius; R1 — radius 1; Rs — radial sector; Ir-m, 2r-m, 3r-m — first, second, and third radio-medial cross-vein; 1r-rs, 2r-rs — first, second radial cross-vein; Sc — subcosta 1 (i.e., free sector of subcosta); Sc+R — subcosta (i.e., fused subcosta proper plus radius); Cells: A — anal cell; C — costal cell; Cul — cubital cell 1; Culb — cubital cell 1b; DA — distal anal cell; IM, 2M, 3M — first, second, and third medial cell; M-Cu — medio-cubital cell; PA — proximal anal cell; R — radial cell; 1R1, 2R1, 3R1 — first, second, and third cell radius 1; 1Rs, 2Rs, 3Rs — first, second, and third radius sector cell. is applied to structures on the ovipositor, which were called pores by Vikberg (1982) and Viitasaari (2002). The membranous lamella along the protibial grooming spur (the anterior spur) is named velum, which is situated on the surface of the spur facing tar- somere | (Schénitzer and Lawitzky 1987). Results Phylogenetic analyses Our focus here is nomenclature and delimitation of genera. Accordingly, we discuss the implications of phylogenetic results only from this perspective. The results largely agree with previous analyses (Nyman et al. 2006; 2010) regarding congruence with current taxonomy: Hemichroa militaris is distinct from other Hemichroa; Stauronematus does not belong to Pristiphora; Pristiphora, Nematus, and some other genera (as delimited in Taeger et al. 2010) in the Euura clade are non-monophyletic; and most species belong to two large clades, Euura and Pristiphora (Figs 2-5). (Note that Dineura virididorsata in Nyman et al. 2006 was a misidentified larva of Nematinus acuminatus; the name has been corrected in GenBank, accessions DQ302173 and DQ302261). The only important difference when it comes to delimitation of genera compared to the previous Marko Prous et al. / Journal of Hymenoptera Research 40; 1-69 (2014) ee __Pistiphoral G6_Mesoneura_opaca N2_Fagineura_crenativora 3b_Craesus_alniastri G4 Craesus_septentrionalis 1p_Nematus_carpini erythrogaster 1/72 3p_Nematus_erythrogaster group 5r_Nematus_abbotii | abbotii 4/81 &DO_Nematus_princeps | group 5f_Nematus_lucidus F6_Nematus_(Paranematus) tulunensis E5 Pristiphora_(Stauronematus) platycerus K6_Cladius_(Priophorus)_compressicomis J7_Cladius_(Trichiocampus)_aeneus JX_Cladius_(Cladius) pectinicornis J5_Hoplocampa_mariatti G3_Hoplocampa_oregonensis H8_Susana_annulata G7_Craterocercus_fraternalis Monocellicampa_pruni_JX566509 1s_Hemichroa_australis J8_Hemichroa_crocea K2_Platycampus _luridiventris DM_Dineura_pullior P1_Hemichroa_militaris H3_Nematinus_acuminatus B5_Nematinus_fuscipennis P2_Fallocampus_americanus H1_Anoplonyx_apicalis J2_Pseudodineura_fuscula 1/86 6r_Pseudodineura_mentiens Pseudodineurini N4_Endophytus_anemones 4r_ Kerita_fidala F7_Caulocampus_acericaulis 1/99 Nematus shigher* Nematinae Cladius @=1/100 1/70 Dineura_ |Dineurini 1/99 Nematinae 1/88 S039_Hoplocampoides_xylostei P4_Phymatocera_aterrima 8) Metallus_pumilus 1/74 ee 4k_Profenusa_thomsoni S080_Periclista_sp U0 $026_Blennocampa_phyllocolpa MX_Endelomyia_aethiops L1_Tenthredo_notha $040_Siobla_ruficornis P8_Strongylogaster_tacita S079 _Nesolandria_morio N9_Eriocampa_ovata S069_Empria_fletcheri 0.2 Figure 2. Phylogenetic tree of Nematinae based on the Bayesian analysis. Numbers above or right of branches show Bayesian posterior probabilities (PP) followed by bootstrap proportions (%, BP) from the corresponding ML analysis. Branches receiving maximum support (PP=1, BP=100%) are denoted by a black dot. Support values for weakly supported branches (PP<0.9 and/or BP<70) are not shown. Euura and Pristiphora clades are collapsed here, but are fully shown in Figs 3 and 4. The inset shows the outline of the full tree including the Euura and Pristiphora clades. Nomenclature is according to Taeger et al. (2010). Voucher ID numbers (e.g. G6) correspond to specimen identifiers in previous phylogenetic trees with different species names. The scale bar shows the number of estimated substitu- tions per nucleotide position. The genera of Nematinae (Hymenoptera, Tenthredinidae) i XXa_Nematus_ferrugineus_group 6b_Nematus_melanocephalus 6m_Nematus_bohemani 1e_Amauronematus_berolinensis 8a_Nematus_distinguendus 6c_Nematus_caeruleocarpus 5m_Nematus_salicis H9 Pachynematus (Larinematus) imperfectus 1/93 p— 2f_Nematus_reticulatus Xd_Nematus_tomeensis 4e Nematus_zebratus J4_Pikonema_dimmockii E7_Pikonema_scutellatum M1_Nematus_miliaris 7f_Nematus_nigricornis Ya_Amauronematus_spiniserra 1/71 JL 9¢ Amauronematus_toeniatus 1/88]L_ 6f Amauronematus_abnormis 4/34] A5_Amauronematus_amplus 4f Amauronematus_krausi 9d_Amauronematus_vittatus_group 1d_Amauronematus_humeralis 4/99 —= 2d_Amauronematus_histrio_group 1n_Amauronematus_nuorbinjargi 8n_Amauronematus_godmani 5e_Amauronematus_histrio_group 1/89 2a_Amauronematus_(Pontopristia) sp A7_Amauronematus_(Pontopristia)_sp 4m_Amauronematus_brunneus @=1/100 0.99/99 C5_Pontania_(Eupontania) norvegica AX _Pontania_(Eupontania)_pustulator 4/97 C7_Pontania_(Eupontania) herbaceae T2_Euura_lanatae 1/90 1/97 CX_Euura_venusta C8_Euura_sp 0.97/87 C3_Pontania_dolichura C2_Phyllocolpa_carinifrons 9n_Phyllocolpa_leucosticta E9 Phyllocolpa_cf__tuberculata D6_Nematus_melanaspis 479 7e_Pachynematus_comiger 9m_Amauronematus_nimbus N6_Amauronematus_(Brachycoluma) sp K8_Nematus_lipovskyi 1785 G5 _Pachynematus _(Polynematus) annulatus 1/93 Xf_Nematus_myosotidis 3a_Nematus_tibialis H5 Pachynematus (Bacconematus) pumilio M3_Pachynematus_(Eitelius) dentatus F1_Pachynematus_(Epicenematus) montanus F5_Nematus_ribesii 4d Nematus_leionotus_group 9b_Nematus_viridissimus 7d_Nematus_spiraeae 7n_Pachynematus_sp Other Nematinae 02 Figure 3. The Euura clade of the Bayesian phylogenetic tree shown in Fig. 2. Numbers above or right of branches show Bayesian posterior probabilities (PP) followed by bootstrap proportions (%, BP) from the corresponding ML analysis. Branches receiving maximum support (PP=1, BP=100%) are denoted by a black dot. Support values for weakly supported branches (PP<0.9 and/or BP<70) are not shown. The inset shows the outline of the full tree. Nomenclature is according to Taeger et al. (2010). Voucher ID numbers (e.g. 6b) correspond to specimen identifiers in previous phylogenetic trees with different species names. The scale bar shows the number of estimated substitutions per nucleotide position. 8 Marko Prous et al. / Journal of Hymenoptera Research 40; 1-69 (2014) F2_Pristiphora_abietina 4/76 1c_Pristiphora_compressa 4c_Pristiphora_pallida J3_Pristiphora_(Sharliphora)_nigella M4 _Pristiphora_alpestris 1/85 3e_Pristiphora_siskiyouensis K7_Pristiphora_erichsonii 3c_Pristiphora_wesmaeli 4a_Pristiphora_bufo Xa_Pristiphora_testacea F3 Pristiohora_coactula Mé6_Pristiphora_geniculata Xe_Pristiphora_maesta 2e Pristiphora_cincta 5d_Pristiphora_thalictri 1/99 6d_Pristiphora_bivittata SSS 1a_Pristiphora_angulata 6a_Pristiphora_leucopus 6n_Pristiphora_nigra Of Pristiphora ‘simulans’ 5a_Pristiphora_retusa H2_Pristiphora_(Micronematus) monogyniae 5c_Pristiphora_biscalis @=1/100 K5_Pristicampus_incisus 193 m_Pristiphora_punctifrons_viridana 1/99 0.94/99 1f_Pristiphora_leucopodia 2b _Pristiphora_nigriceps 1/86 1b Pristiphora_laricis D9_Pristiphora_mollis L7_Melastola_ferruginosa 0.92/76 J1_Pristola_macnabi L9_Neopareophora_litura 3d_Nepionema_helveticum Other Nematinae 0.2 Figure 4. The Pristiphora clade of the Bayesian phylogenetic tree shown in Fig. 2. Numbers above or to the right of branches show Bayesian posterior probabilities (PP) followed by bootstrap proportions (%, BP) from the corresponding ML analysis. Branches receiving maximum support (PP=1, BP=100%) are denoted by a black dot. Support values for weakly supported branches (PP<0.9 and/or BP<70) are not shown. ‘The inset shows the outline of the full tree. Nomenclature is according to Taeger et al. (2010). Voucher ID numbers (e.g. F2) correspond to specimen identifiers in previous phylogenetic trees with dif- ferent species names. The scale bar shows the number of estimated substitutions per nucleotide position. analyses is the phylogenetic position of the type species of Nematus, N. lucidus. The addition of the nuclear NaK gene to the dataset causes NV. /ucidus, as well as Mesoneura, Fagineura, Craesus, and some other species of Nematus to move away from the Euura clade (placed closer in Nyman et al. 2006; 2010) and approach the Pristiphora clade (Figs 2 and 5). Considering previous phylogenetic results and the results reported here (Figs 2-5), we propose in the Taxonomy section several changes compared to Taeger et al. (2010). Advantages and disadvantages of this and alternative taxonomies are high- lighted in the Discussion section. The genera of Nematinae (Hymenoptera, Tenthredinidae) 9 BC_ZSM_HYM_06891_Epicenematus_pallescens H5_Bacconematus_pumilio M3_Eitelius_dentatus G5_Polynematus_annulatus BIOUG01862-E10_Pteronidea_ventralis F5_DEIGISHym17040_Kontuniemiana_ribesii BC_ZSM_HYM_11332_Holcocneme_vicinus T4_Euura_mucronata evn YX_Cryptocampus_amerinae 82/1 R5_DEIGISHym11810_Eupontania_vesicator Q5 Pontania_proxima Euura 80/1 E9_Tubpontania_cf._anomaloptera X9_Phyllocolpa_leucapsis J4_Pikonema_dimmockii 10.98 2f_Lindqvistia_reticulatus —_——— DEIGisHym4855_Amauronematus_stenogaster A7_Pontopristia_cf.__amentorum 87/4 H9_DEIGISHym11466_Larinematus_imperfectus 7n_BC_ZSM_HYM_16651_Pachynematus_cf._clitellatus H6_Brachycoluma_viduata DEIGISHym19657_Decanematus_malaisei J3_DEIGISHym11927_Sharliphora_nigella 84/1 __F2 BC_ZSM_HYM_09367_Lygaeonematus_abietinus sori | Xa_DEIGISHym19642_Pristiphora_testacea F3_Lygaeotus_coactula K9_Sala_chlorea H2_DEIGISHym11324_Micronematus_monogyniae oat Ls 8t_Gymnonychus_abbreviatus Pristiphora K5_Pristicampus_incisus 70/0.94 DEIGISHym10991_Pristicampus_arcticus rit J1_Pristola_macnabi 3d_Nepionema_helveticum a 50/0.91 L9_Neopareophora_litura 7 1b_DEIGISHym11555_Oligonematus_laricis 3r_Diphadnus_appendiculatus G6_Mesoneura_opaca N2_Fagineura_crenativora DEIGISHym17938_Paranematus_wahlbergi_wahlbergi F6_DEIGISHym19679_Paranematus_tulunensis 5r_Hypolaepus_abbotii Nematus ‘G4_Craesus_septentrionalis 5f_Nematus_lucidus DEIGISHym15246_Craterocercus_obtusus 7_Craterocercus_fratemalis — _ DEIGISHym15241_Adelomos_cleone E5_Stauronematus_platycerus 1s_Hemichroa_australis K2_BC_ZSM_HYM_11000_Platycampus_luridiventris P1_Vama_militaris DEIGISHym20368_Dineura_virididorsata B5_BC_ZSM_HYM_15192_Nematinus_fuscipennis. P2_Fallocampus_americanus Monocellicampa_pruni_JX566509 DEIGISHym11436_Pseudodineura_parvula J2_Phyllopais_fusculus N4_Endophytus_anemones 4r_DEIGISHym15265_Kerita_fidala F7_Caulocampus_acericaulis JX_DEIGISHym19475_Cladius_pectinicornis K6_BC_ZSM_HYM_16700_Priophorus_compressicornis BC_ZSM_HYM_03232_Trichiocampus_grandis DEIGISHym17823_Moricella_rufonota ~ DEIGISHym11103_Hoplocampa_brevis G3_Macgillivrayella_oregonensis K1_Susana_cupressi H8_Susana_annulata @= 100/1 01 Figure 5. Phylogenetic relationships of type species for which at least 400 bp of the Col barcode region was available. The tree was reconstructed according to a ML analysis allowing a separate GTR+I+G4 model of substitution for each gene. Numbers above or right of branches show bootstrap proportions (%, BP) fol- lowed by posterior probabilities (PP) from the corresponding Bayesian analysis. Branches receiving maxi- mum support (BP=100%, PP=1) are denoted by a black dot. Support values for weakly supported branches (PP<0.9 and BP<50) are not shown. Nematinae was not monophyletic in the Bayesian analysis, because Moricella rufonota (only 658 bp of Col available) was weakly placed as sister to the Strongylogaster+ Nesoselan dria clade. Some terminals in the dataset are composites of two conspecific specimens (indicated by two ID numbers, for example J3 and DEIGISHym11927 or F2 and BC ZSM HYM 09367). Four non-type spe- cies (Pristicampus incisus, Paranematus tulunensis, Craterocercus fraternalis, and Susana annulata) were also included in the analysis, because the Col or NaK (for S. cupressi) sequences available for the type species of the respective genera were not sufficient to place them reliably in the tree. Type species in bold are the basis for genera defined here. Outgroup taxa have been collapsed for simplicity. Voucher ID numbers (e.g. F2) correspond to specimen identifiers in previous phylogenetic trees with different species names. ‘The scale bar shows the number of estimated substitutions per nucleotide position. 10 Marko Prous et al. / Journal of Hymenoptera Research 40; 1-69 (2014) Key For some smaller genera and species groups there are comprehensive recent keys to species available, which we have referred to in the key. Because the morphological separation of Euura, Pristiphora, and Nematus can be difficult (see Discussion), there are two places (couplets 15 and 17) in the key where we have not separated them completely. Instead, we have given short descriptions of a few minor groups which run together with the genus intended to be keyed out. Full resolution of these prob- lems requires species-level revisions. The key is arranged in alternating pages of text and plates. The couplets are illustrated by a plate on the facing page. We recommend using the key with two pages side by side, so that couplets and corresponding figures are simultaneously visible. 1 a Fore wing shortened, apex usually not reaching tip of abdomen, veins often strongly aberrant. [Some arctic or high elevation specimens of Euura and Pristiphora, e.g. females ol: Buura abnormis TOlmeren, 1883) | cccvvetnecpiiie users omen aaiesisvene ode 10 — aa Fore wing normal, apex extending beyond tip of abdomen, veins rani lly cev elo Peds. ore. exe eseevyceeal eeprecaath yn are wergleeseevocpeaien were 2 2(1) a Vein 21r-m present, joining vein M proximal of 2m-cu...... ee 43 — aa Vein 21-m joining vein M either distal of 2m-cu or very slightly proximal of 2m-cu (few aberrant specimens), or vein 2r-m absent.............ee 3 a Vein 2A of hind wing complete, cell A closed. LiYPaleai biChakiat Vet oid oer aeeal Baw eon Wen aU TRAE UNS MP AMR TMA gen mee 4 = aa Vein 2A of hind wing incomplete, cell A open distally. bb Body length 2-6 mm. [Psendodineurint antdutéwy Pristi prior.) x. iceactoot ctma rent epbetss te pneesibsetedonrons ion 40 4(3) a _ Base of vein 2A+3A complete and curved up to 1A, cell PA closed .. 33 = aa_ Base of vein 2A+3A incomplete and straight, cell PA open distally..... 5 5(4) em Velin bts ProCite, Me, Ret Sextus rsa nama dled | Me aay de orate ance 23 _ AED UPA TTS SCO ok ac vas a etal et et nar edt ea aes et beat nan altSabaleatietaeeeee 6 6(5) a Left mandible markedly constricted near middle and right mandible fapeted repularly LOWAlds Apes. suche. es. outonnasttn sate oneieleerasenantioe’s 7 — aa Left and right mandible both tapered regularly towards apex........... 18 The genera of Nematinae (Hymenoptera, Tenthredinidae) 11 la Euura abnormis, female laa E. abnormis, male (1a and laa from Benson 1958) 2a Hoplo- campa chrysorrhoea 2aa Nematus lucidus 3a Hoplocampa chrysorrhoea 3aa Pseudodineura enslini 4a Platycampus luridiventris 4aa Nematus lucidus 5a Hemichroa australis 8aa Nematus lucidus 6a Dinematus krausi 6aa Nematinus fuscipennis (left), Dineura virididorsata. 12 7 (6) -*) bb cc Marko Prous et al. / Journal of Hymenoptera Research 40; 1-69 (2014) Tarsal claw without basal lobe and subapical tooth absent or shorter than apical tooth. Clypeus deeply emarginate, subtruncate, or truncate. Apex of flagellomeres in males not produced ventrally... 8 Tarsal claw with basal lobe and subapical tooth present, erect, well separated from apical tooth, and longer than apical tooth. Clypeus not deeply emarginate. Apex of flagellomeres in male slightly produced ventrally. Two species. Key to species by Liston (2007). Palaearctic. Fe ere Ln Seine See My Tere ae Stauronematus Benson, 1953 Notauli sharply outlined. Exposed part of membrane between mesoscutellar appendage and postnotum of mesothorax anteriorly widened or not «0.00... eeeeeeeeeeees 9 Notauli hardly outlined. Exposed part of membrane between mesoscutellar appendage and postnotum of mesothorax anteriorly widened. Four species of the P resinicolor and P macnabi groups. Formerly Pristola and Melastola. Key to species by Wong (1968). Nearctic. Baengttt Mant ae hie Protea sauce avert ee etna Pristiphora Latreille, 1810 in part The genera of Nematinae (Hymenoptera, Tenthredinidae) 13 t 8b/8bb 7a From left to right: Euura pumilio, E. clitellata, Nematus lucidus, E. ribesii Taa Stauronematus platycerus Tb N. septentrionalis Tblbb Pristiphora mollis (upper right), P testacea Tbb S. platycerus 7c N. lucidus Tc S. platycerus 8a N. lucidus 8b N. lucidus 8aa P macnabi 8bIbb P. resinicolor. 14 9(8) 10(1, 9) op bb cc op Marko Prous et al. / Journal of Hymenoptera Research 40: 1-69 (2014) Cell 1Rs and 2Rs fused, vein 2r-m absent. Clypeus emarginate. Tarsal claw with long subapical tooth. About 60 species of Euura. Former Euura s.str., Alpinematus, and some Pontania. Holarctic. neath ee Unt aat ste Nader «pe patee Sannaesh Sateen rnd Suhel Euura Newman, 1837 in part Cell 1Rs and 2Rs not fused, vein 2r-m present. Clypeus emarginate or truncate. Tarsal claw without or with short or long subapical tooth................ 10 Clypeus clearly and rather deeply emarginate, and tarsal claw with long subapical tooth... eee eeeeseeseeeeeeeeee 11 Clypeus more or less truncate, or/and tarsal claw without or with small subapical tooth .................. 16 The genera of Nematinae (Hymenoptera, Tenthredinidae) 15 . TRS + 2Rs oe es ry { 9a Euura mucronata Yaa Nematus lucidus 9bI9bb E. mucronata 9cI9cC E. mucronata 9bb Pristiphora testacea 9cC E. clitellata \0a N. septentrionalis (left), N. lucidus \Oaa P mollis (left), P testacea \0b N. lucidus \Obb E. pumilio (left), E. clitellata. 16 11(10) 12(11) 13(12) a” cc dd -¥) bb cc Marko Prous et al. / Journal of Hymenoptera Research 40; 1-69 (2014) Height of eye in lateral view about 2—3 times as long as distance from dorsal margin of eye to dorsalmost point of head. Sawsheath emarginate in dorsal view. Tangium of lancet with campaniform sensilla (“pores”). Valviceps not divided into pseudoceps and paravalva, without valvispina. Small, 4.0-5.5 mm, black and yellow-brown; thorax (except pronotum), abdomen above, head (almost entirely) and antennae black; clypeus, labrum and base of mandibles white. In the forewing costa same colour as pterostigma (usually pale). Three species of the P arctica group. Formerly Pristicampus. Revision by Zinovjev (1993). Palaearctic. as LE ia tot corti se ce le AL cad Pristiphora Latreille, 1810 in part Height of eye in lateral view usually about 3—4 times as long as distance from dorsal margin of eye to dorsalmost point of head. Sawsheath usually not emarginate in dorsal view. Tangium of lancet without campaniform sensilla, if present, then aa apply. Valviceps divided into pseudoceps and paravalva, often with valvispina... 12 Metatarsomere | 2.0—3.0 times as wide as width of metatarsomere 2. 22 species of the NV. septentrionalis group. Formerly Craesus. Holarctic, Oriental. ebay ohn ents a heartgdey eal eng tae ag tiatea eee GeateLsastes Nematus Panzer, 1801 in part Metatarsomere | 1.0—1.5 times as wide as width of metatarsomere 2...13 Metatarsomere 1 with long and deep groove on anterior and posterior surfaces. Valvispina widened preapically. Body length 8-12 mm. About four species of the NV. abbotii group, including N. princeps Zaddach, 1876. Key to Nearctic species by Smith (2008). Holarctic. Edi stecs otaeetasta Buk eshte sanded gsh ueredesQbede etal eahiles Nematus Panzer, 1801 in part Metatarsomere | at most with short and shallow groove on anterior and posterior surfaces. Valvispina evenly tapering towards tip. Body length Sal tani acti. ones cesneghven ce ees patensshavectipavesslteasnesetiewnsastens 14 The genera of Nematinae (Hymenoptera, Tenthredinidae) Ly 11cc pseudoceps paravalva —- valvispina. lla Pristiphora arctica \laa Euura reticulata \\b P incisa \\bb E. reticulata \\¢ P arctica \\ ce E. re- ticulata \\d P arctica \\dd E. vicina \2a Nematus septentrionalis \2aa E. caeruleocarpus \3a N. abbotii I3aa E. vicina (left), E. caeruleocarpus \3b N. princeps \3bb E. vicina. 13b 18 14(13) 15(14) a Marko Prous et al. / Journal of Hymenoptera Research 40: 1-69 (2014) Mesothoracic katepimeron extensively covered with hairs. Sawsheath slightly emarginate or not emarginate in dorsal view. Tangium of lancet with campaniform sensilla (see Fig. 1 1c). Sawsheath slightly emarginate in dorsal view in the type species, F crenativora Vikberg & Zinovjev, 2000, not emarginate in F quercivora Togashi, 2006. Two species. Key to species by Togashi (2006). East Palaearctic. Rr Rede rer wt Fagineura Vikberg and Zinovjev, 2000 in part Mesothoracic katepimeron with at most few hairs. Sawsheath usually not emarginate in dorsal view. Tangium of lancet without campaniform sensilla (see Fig. 1 1cc)...... 15 At least terga 2—3 or at most terga 1-6 reddish. Pronotal angles reddish. Stigma dark. Legs largely red; coxae, trochanters and hind tarsi black. Body elongate, torpedo-shaped. The type species of Nematus, N. lucidus Panzer, 1801. 7—11 mm. Palaearctic. spec Beare eee ds ene gh ewe degeah alte ny antigen heels Nematus Panzer, 1801 in part aa—ee Combination of characters not as in a—e. Large part of about 700 Euura species, e.g. most of former Amauronematus, Nematus (Pteronidea), Phyllocolpa, and Pontania. Holarctic, Oriental, and introduced to Afrotropic, Neotropic and Australasia. The type species, E. mucronata, keys out in couplet 9 or 22. Pocgales hash We nde AU AG Seas ite MO ag tsic Euura Newman, 1837 in part Body 8 mm; head and antennae entirely black; thorax dorsally mostly reddish, ventrally mostly black; pronotum and tegulae yellowish; stigma dark brown; legs black; abdomen yellowish, sawsheath black; mesepisternum smooth. One species, NV. noblecourti Lacourt, 2006. Nematus? West Palaearctic. NeseeR rie eat ecct val tala bars oeecchnosdleie soa bum enieaeedrcbere Nescianeura Lacourt, 2006 Body 5.5-10.5 mm, black; pronotal angles and tegulae yellowish; stigma dark brown; legs largely pale; hind tibia at least in basal third pale; mesepisternum smooth; sawsheath in dorsal view narrowing towards the apex, apically broadly rounded. On Lonicera. About six species of the NV. wahlbergi group. Formerly Paranematus. Keys to species by Vikberg (1972), Zinovjev (1978). Palaearctic. sgh Ue casacn dle RAT Bd dope avinsva ins Wives dies ke Nematus Panzer, 1801 in part Body 4.5—7.0 mm, black or with yellowish or reddish abdomen, sometimes also thorax largely reddish; stigma dark brown; hind tibia slightly widened with an indistinct longitudinal groove; metatarsomere | cylindrical, without a groove; sawsheath short and rounded in lateral view, wide in dorsal view; cerci longer than sheath; lancet with lateral spines on annuli; male penis valve is slightly curved, with stout valvispina, and a rather low, rounded adjacent lobe. About six species of the LV. erythrogaster group, including N. /ucens (Enslin, 1918) and N. umbratus Thomson, 1871. Keys to species by Liston et al. (2006), Smith (2008). Holarctic. RANI fete con BNR a Aaron in dal ein fate nt ena Nematus Panzer, 1801 in part The genera of Nematinae (Hymenoptera, Tenthredinidae) 15a,b,c,e 14a, 14b Fagineura crenativora \4aa,l4bb Euura reticulata \5a,b,c,e,d Nematus lucidus. 20 16(10) a0 Ff Marko Prous et al. / Journal of Hymenoptera Research 40: 1-69 (2014) Head length behind eyes 0.2—0.4 times as long as eye length in dorsal view. Sawsheath frequently distinctly emarginate in dorsal view. Clypeus more or less truncate (see Fig. 10aa). Apex of vein C usually swollen, at the point of origin of vein Rs+M from R cell C is usually only about as wide as vein R. Medial surface of head near antennal sockets markedly elevated in lateral view and angular below medial pit. In female, apical part of abdomen sometimes laterally compressed, so that it appears narrow in dorsal view. Tangium of lancet with campaniform sensilla. In male, tergum 8 without long medial projection, if present, then a, c, d, and e apply. About 240 species, Holarctic, Oriental, and Neotropical. nites Pisani tate sal PN he eles Caan seth Pasa Pristiphora Latreille, 1810 in part Body 7 mm; head (including antennae), thorax, and abdomen dorsally mostly black, ventrally yellowish, except ventral black half of mesepisternum; pronotum, tegulae, and stigma yellowish, legs mostly yellowish; mesepisternum smooth. One species, D. krausi Lacourt, 2006. Pristiphora? West Palaearctic. a eet CUD COATT IO ANU DP Dinematus Lacourt, 2006 in part Head length behind eyes 0.4—0.7 times as long as eye length in dorsal view. Sawsheath usually not distinctly emarginate in dorsal view. Clypeus usually deeply emarginate (see Fig. 10a). Apex of vein C usually less swollen, at the point of origin of vein Rs+M from R cell C is about twice as wide as vein R or wider. Medial surface of head near antennal sockets sometimes elevated in lateral view and slightly or not angular below median pit. In female, apical part of abdomen not laterally compressed. Tangium of lancet usually without campaniform sensilla. In male, tergum 8 usually with long medial projection... 17 The genera of Nematinae (Hymenoptera, Tenthredinidae) au a to ca 16a Pristiphora pallidiventris \6aa Euura annulata \6b P pallidiventris \6bb E. annulata 16d P pallidiventris \6dd E. annulata \6e P. testacea \6ee E. stenogaster 16 P pseudodecipiens | 6ff E. stenogaster 16g P compressa \6gg E. annulata \ 6h P testacea \6hh E. imperfecta (left), E. clitellata. 22 17(16) 18(6) -¥) ao” cc dd op bb cc Marko Prous et al. / Journal of Hymenoptera Research 40; 1-69 (2014) Height of eye in lateral view about 2—3 times as long as distance from dorsal margin of eye to dorsalmost point of head, and sawsheath emarginate in dorsal view. Tangium of lancet with campaniform sensilla. Valviceps not divided into pseudoceps and paravalva, without valvispina. Small, 4.0-5.5 mm, black and yellowish brown; thorax (except pronotum), abdomen above, head (almost entirely) and antennae black; clypeus, labrum and base of mandibles white. Forewing costa same colour as pterostigma (usually pale). Three species of the P arctica group. Formerly Pristicampus. Revision by Zinovjev (1993). Palaearctic. He LE det coer ete te SO Set, AL ace Pristiphora Latreille, 1810 in part Height of eye in lateral view about 3-4 times as long as distance from dorsal margin of eye to dorsalmost point of head, or/and sawsheath not emarginate in dorsal view. Tangium of lancet usually without campaniform sensilla. Valviceps divided into pseudoceps and paravalva, often with valvispina. Part of about 700 Euura species, e.g. most of former Pachynematus. Holarctic, Oriental, and introduced to Afrotropic, Neotropic and Australasia. 5 dasintcc Osan sabtoit leans Meagan Saas emaenasumenn cp aRatn a Euura Newman, 1837 in part Body 5.5—10.5 mm, black; stigma dark brown; pronotal angles and tegulae yellowish; legs largely pale; hind tibia at least in basal third pale; claws with large subapical tooth; mesepisternum smooth; sawsheath in dorsal view narrowing towards the apex, apically broadly rounded. On Lonicera. About six species of the NV. wahlbergi group. Formerly Paranematus. Keys to species by Vikberg (1972), Zinovjev (1978). Palaearctic. Spree Sashes olde ale caves eee Seale cots chchatte oe te sale Nematus Panzer, 1801 in part Body 6.0-7.5 mm, black; tegulae, pronotal angles, stigma, tarsi and tibiae yellow, femora usually only partly yellow; claws with small subapical tooth, mesepisternum rough. One species of Pristiphora, P mollis (Hartig, 1837). West Palaearctic, Nearctic. ee a SA ee ORR Ronee Ce RE Pristiphora Latreille, 1810 in part Tarsal claw with subapical tooth larger than apical tooth. Malar space 0.1—0.6 times as long as diameter of front ocellus. Clypeus truncate or slightly and narrowly emarginate. Few specimens of Moricella. Oriental. RP AR TT ae RR, SR I en ea Moricella Rohwer, 1916 in part Tarsal claw with subapical tooth smaller than apical tooth or absent. Malar space 1.0—2.0 times as long as diameter of front ocellus. CTs PeUs WH Oly CH AT OATS sa undies lauerineaan codes eoat aneaeraacibasatsnapehateat 19 The genera of Nematinae (Hymenoptera, Tenthredinidae) 23 pseudoceps paravalva valvispina ; cc 18b/18¢c he eee : 18bb/18cc 17a Pristiphora incisa \Taa Euura imperfecta \7Tb P incisa \Tbb E. annulata \Tc P arctica I7cc £. annulata 11d P arctica \1dd E. vicina 18a Moricella rufonota \8aa Nematinus fusci- pennis (left), Dineura virididorsata \8bI1 8c M. rufonota \8bb/18cc N. fuscipennis. 20(19) 21(20) -¥) cc dd Marko Prous et al. / Journal of Hymenoptera Research 40: 1-69 (2014) Anterior protibial spur with velum. Vein Sc usually situated before point of origin of vein M from R..... 20 Anterior protibial spur without velum, but with hairs. Vein Sc usually situated beyond or at point of origin of vein M from R. Ten species, including D. militaris (Cresson, 1880) (previously in Hemichroa). Holarctic and Oriental. pede 1 ies eit, 3nd RN A ee 3 I Dineura Dahlbom, 1835 in part Tarsal-claveswithesubapical tootht.00 2 .n.2 ni) ee teh ee 21 Tarsal claw without subapical tooth. Only few Anoplonyx specimens key out here. Holarctic. Brac a cece MR ted 2 A FcR Mie Anoplonyx Marlatt, 1896 in part Abdominal tergum IX in female in lateral view more than 3 times as long as tergum VUI. Penis valve in male of several species with filament. Anterior depressed section of metepisternum along metepimero- episternal suture 0.3—0.7 times as wide as posterior section. Cell 1Rs and 2Rs not fused because vein 2r-m present (see Fig. 9aa). 26 species. Holarctic. sghacneste Mes seensns Fults aie Mace k? AR: eon thilieca hut A Rat! Nematinus Rohwer, 1911 Abdominal tergum IX in lateral view in female less than 2 times as long as tergum VIII. Penis valve in male without filament. Anterior depressed section of metepisternum along metepimero- episternal suture 0.1—0.3 times as wide as posterior section. Cell 1Rs and 2Rs often fused because vein 2r-m absent (see Fig. 9a) ....22 The genera of Nematinae (Hymenoptera, Tenthredinidae) 25) a 19a Nematinus fuscipennis (left), N. bilineatus \Yaa Dineura militaris \9b N. fuscipennis \9bb D. viri- didorsata 20a N. fuscipennis (left), E. clitellata aa Anoplonyx ovatus 2a N. fuscipennis laa Euura vesicator 21 b N. fuscipennis (left), N. bilineatus 2\ bb E. vesicator 2\¢ N. fuscipennis 2\ cc E. vesicator. 22(21) 23(5) 24(23) bb cc -¥) ceore cc Marko Prous et al. / Journal of Hymenoptera Research 40; 1-69 (2014) Katepimeron extensively covered with hairs. Sawsheath slightly emarginate or not emarginate in dorsal view. Tangium of lancet with campaniform sensilla. Sawsheath slightly emarginate in dorsal view in the type species, F crenativora Vikberg & Zinovjev, 2000, not emarginate in F quercivora Togashi, 2006. Two species. Key to species by Togashi (2006). East Palaearctic. i chuitdafbeied Maine iaaavensbinnals Fagineura Vikberg and Zinovjev, 2000 in part Katepimeron without hairs. Sawsheath not emarginate in dorsal view. Tangium of lancet without campaniform sensilla. About 150 species of Euura, including former Euura s.str., Pontania. Holarctic, Oriental, and introduced to Neotropic and Australasia. sdheeanialintettes diesel nc ea lena banned denies duaehes aaannsatethet Euura Newman, 1837 in part Petiole of anal cell 1A shorter than CU-a ........ceeeececeeeeeeccceecccceuseeeeceees 24 Petiole.of anial*celll LA TOm@Er THAN Clica ticks eiaesdsenseronapenadouedenselpeatiss 26 Antenna about 2 times as long as width of head. Left (illustrated) and right mandible both tapered regularly towards apex. Body Lene tao mails sssh te teoauceotardtniaca tease taeedanteetanctvaiealtt Sedna 25 Antenna about 4 times as long as width of head. Left mandible constricted near middle and right mandible tapered regularly towards apex. Body length 10-15 mm. Four species. Key to species by Wei and Nie (2008). East Palaearctic and Oriental. ate ® Peet, er nee eee Pes A Pae ere sks Ae Megadineura Malaise, 1931 The genera of Nematinae (Hymenoptera, Tenthredinidae) 2H a +s “i 22 — 23aa 22a,22b Fagineura crenativora 22aa,22bb Euura vesicator 23a Megadineura grandis 23aa Di- neura virididorsata 24a Moricella rufonota 24aa M. grandis 24b M. rufonota 24bb M. grandis. 28 25(24) 26(23) 27 (26) op bb cc op bb cc Marko Prous et al. / Journal of Hymenoptera Research 40: 1-69 (2014) Clypeus truncate or slightly and narrowly emarginate. Anterior protibial spur without velum, but with hairs. Sawsheath emarginate in dorsal view. Three species. Oriental. vr tes et Se oral reo re hrc Moricella Rohwer, 1916 in part Clypeus deeply and widely emarginate. Anterior protibial spur with velum. Sawsheath not emarginate in dorsal view. One species, K. planaritibia Togashi, 1964. East Palaearctic. saeedlc Pa chadhd oar ener setotach tet hctes bal vesteaai duets Neaiseasl es t&h Katsujia logashi, 1964 Clypeus long, width 2.2—2.6 times as long as length. Labrum apically emarginate. Antenna 1.4—2.0 times as long as width of head. Left and right mandible both tapered regularly towards apex........... 27 Clypeus short, width 2.8—4.0 times as long as length. Labrum apically rounded. Antenna 1.4—4.0 times as long as width of head. Left and right mandible both tapered regularly towards apex or left mandible markedly constricted near middle and right mandible tapered LeSularly: tOWalds APEK convmaccreycosmayvenr doconmvecsinayasuenesenn doccomneceteiiee 28 Anterior protibial spur without velum. Outer margin of eye without distinct furrow. Clypeus deeply emarginate. One species, A. cleone Ross, 1935. Only the type specimen is known to have a vein r (cross-vein of cell R1) of hind wing present. Nearctic. fae ich Ti a de 5 Ae cred air en nik Meas hn Adelomos Ross, 1935 Anterior protibial spur with velum. Outer margin of eye with distinct furrow. Clypeus slightly emarginate. One species, NV. arquata (Klug, 1816). West Palaearctic. cdapepAdeetAtcbead Jaan iu oagentdanyaha sngnddsterdacetan uhh: Neodineura Taeger, 1989 The genera of Nematinae (Hymenoptera, Tenthredinidae) 29 eb. 4 ( unl. (27ec) 26a/26b 25a Moricella rufonota 25aa Katsujia planaritibia 25b M. rufonota 25bb K. planaritibia 25c M. rufonota 25cc K. planaritibia 26al26b (27cc) Neodineura arquata 26al26b (27c) Adelomos cleone 26aal26bb Dineura virididorsata (left), Mesoneura opaca 27a A. cleone 2Taa N. arquata 21b A. cleone 27bb N. arquata. 28(26) 29(28) 30(29) a0 & Bf cc dd bb cc Marko Prous et al. / Journal of Hymenoptera Research 40: 1-69 (2014) Left mandible markedly constricted near middle and right mandible LACT CG MEO UAT IY LO WALASFAD OI Tuan Lauter suo sRittas aah eeltea eat tala 29 Left and right mandible both tapered regularly towards apex........... 32 Shortest distance between eyes about 2.0—2.5 times as long as height of eye. Antenna 3.5—4.0 times as long as width of head. Malar space 1.5—2.0 times as long as diameter of front ocellus. Clypeus shallowly emarginate. One species, R. fumosa Wei and Nie, 1998. East Palaearctic. Eee (0 RR ern NP EAE AOD Renonerva Wei and Nie, 1998 Shortest distance between eyes 1.1—2.0 times as long as height of eye. Antenna 1.4—3.2 times as long as width of head. Malar space 0.2—2.0 times as long as diameter of front ocellus. Elypeus shallowly ordeeplyeMarcinareis. cc. rrscersrotamsuocaceactesdicrenater 30 Malar space 0.2—0.5 times as long as diameter of front ocellus. Antenna 1.4—1.9 times as long as width of head. Veins 2r-rs, 2r-m, and 2m-cu nearly forming single straight line in anterio-posterior direction. Nine species. Key to species by Wei et al. (2013a). Palaearctic. vg eailvee- ees ch ucnedvncaye wnat dievnlacasue desi tedsiies cateyenns Mesoneura Hartig, 1837 Malar space 0.7—2.0 times as long as diameter of front ocellus. Antenna 1.8—3.2 times as long as width of head. Veins 21-rs, 2r-m, and 2m-cu forming three separate lines in anterio- PUSPEEIOR GE CUIG DL cre 0B 8 a tea chan atte ced etre Otten ra rn ea ach 31 The genera of Nematinae (Hymenoptera, Tenthredinidae) | 30cc 28a Dinematus krausi 28aa Moricella rufonota 29al29c,ccl29d,dd Renonerva fumosa (original photo by Gengyun Niu) 29aa/29cc/29dd Mesoneura opaca 29b R. fumosa (photo by Shaobing Zhang; http://www.sawfly.cn/yftk/ShowPhoto.asp?PhotoID=143) 30a M. opaca 30aa Pristiphora incisa 30b M. opaca 30bb P. arctica 30€ M. opaca 30cc P litura. 32(28) 33(4) bb cc op bb cc -¥) bb cc Marko Prous et al. / Journal of Hymenoptera Research 40: 1-69 (2014) Clypeus truncate or slightly emarginate. Apex of vein C usually swollen, at the point of origin of vein Rs+M from R cell C is usually only about as wide as vein R. Sawsheath slightly emarginate in dorsal view and/or tarsal claw without subapical tooth. Four species of Pristiphora. Formerly Nepionema, Neopareophora. Nearctic, West Palaearctic. Fey ee RU Me Dae rn, PN Pr ee Pristiphora Latreille, 1810 in part Body 7 mm; head (including antennae), thorax, and abdomen dorsally mostly black, ventrally yellowish, except black ventral half of mesepisternum; pronotum, tegulae, and stigma yellowish, legs mostly yellowish; mesepisternum smooth. One species, D. krausi Lacourt, 2006. Pristiphora? West Palaearctic. Nth RSet hate Nlan ce Latent Sen eA Beda selon as Dinematus Lacourt, 2006 in part Clypeus widely and deeply emarginate. Apex of vein C less swollen, at the point of origin of vein Rs+M from R cell C is about twice as wide as vein R or wider. Sawsheath not emarginate in dorsal view and tarsal claw with long subapical tooth. One species of Euura, some females of E. radialis (Smith, 1994), comb. n. Nearctic. PANO: aR Rtas ONT 1, SERRA TERROR VR RT Euura Newman, 1837 in part Tarsal claw with subapical tooth as long as or slightly longer than apical tooth. Malar space 0.1—0.6 times as long as diameter of front ocellus. Clypeus truncate or slightly and narrowly emarginate. Three species. Oriental. Pesach sis: ute td oi Re east ME teeta Moricella Rohwer, 1916 in part Tarsal claw without subapical tooth or with subapical tooth shorter than apical tooth. Malar space 1.0—2.0 times as long as diameter of front ocellus. Clypeus deeply and widely emarginate. Ten species. Holarctic and Oriental. Ec the 15 inh Lisi a Ath Ae RE Dineura Dahlbom, 1835 in part Length of vein R between junctions with veins M and Rs+M longer than first sector of Rs. Male and female flagellum similar: thread or seta-like, with short pubescence. Weil, 218s preselit OGADSENC Spee te are neces na 34 Length of vein R between junctions with veins M and Rs+M not longer than first sector of Rs. Male and female flagellum dissimilar: seta-like, with short pubescence in female, and seta-, comb-, or saw-like, with long pubescence in male. Vein 2r-rs absent. Few aberrant specimens of Cladius. acoder ep eAcbucphattsey tile ouitogiachucytbni ey thy eees Cladius \lliger, 1807 in part The genera of Nematinae (Hymenoptera, Tenthredinidae) 33 3 1a Dinematus krausi (left), Pristiphora litura 3 \aa Euura radialis 3\b P. pallidiventris 3 \bb Nema- tus lucidus 3\c P. litura sawsheath (left), P. helvetica tarsal claw 31. cc (3 1c) E. radialis sawsheath and tarsal claw 32a Moricella rufonota 32aa Dineura virididorsata 32b/32¢ M. rufonota 32bb/32cc D. virididorsata 33a Platycampus luridiventris 33aa Cladius compressicornis 33b Platycampus luridiven- tris 3, 2 33bb C. grandis 3, P (top); C. compressicornis 3, 2 (bottom); C. pectinicornis 3, & (right). 34 34(33) 35(34) 36(35) 37 (36) ono & ® op Marko Prous et al. / Journal of Hymenoptera Research 40: 1-69 (2014) Clypeus widely and deeply emarginate. Malar space clearly shorter than diameter of front ocellus. Anterior half of mesepimeron partly or completely covered with setae. Vein 2r-rs of fore wing present (see Fig. 5a). Nearctic. Six species. Key to most of the species by Smith (1969a). Ae De SARI eed AN REM bE en 2 Craterocercus Rohwer, 1911 Clypeus narrowly and deeply or shallowly emarginate. Malar space equal to or longer than diameter of front ocellus. Anterior half of mesepimeron without setae. Vein 2r-rs of fore wing absent or present (see Figs 5a and 5aa). PACAEC Listy ees ects Pee et oy Pore ae a AE ae etn, ae ee Reo Pee et cae 35 Tarsal claw with subapical tooth. Clypeus 2—3 times as wide as long. Cercus in female 3—20 times as long as Wide... eeeeeeeseeeeeseeees 36 Tarsal claw without subapical tooth. Clypeus 3 times as wide as long. Cercus in female 2—4 times as long as wide. Twelve species. Holarctic. Be ee ee Anoplonyx Marlatt, 1896 in part Distance between pulvilli of metatarsomeres 1 and 2 about twice or more their length. Vein 2r-rs of fore wing absent or present (see Figs 5a and 5aa)......... 37 Distance between pulvilli of metatarsomeres 1 and 2 subequal to their length. Vein 2r-rs of fore wing absent. Two species. Key to species by Smith (1976a) under Platycampus. Nearctic. Seopa, eters Eas me nied ree MWe RA ate Fallocampus Wong, 1977 Anterior protibial spur without velum, but with hairs. Vein 2r-rs of fore wing absent (see Fig. 5aa) 0... eeeeeseseeseeseeeeeeeeees 38 Anterior protibial spur with velum. Vein 2r-rs of fore wing absent Or Present..........sceeceeeseeseeeseeseeeseeeees 39 The genera of Nematinae (Hymenoptera, Tenthredinidae) 35 ‘eal x 34al34b Craterocercus obtusus 34aal34bb Hemichroa australis (left), Anoplonyx apicalis 34¢ C. obtusus 34cc A. apicalis 35a,35b,35c H. australis 35aa,35bb,35cc A. ovatus 36a Platycam- pus luridiventris 36aa Fallocampus americanus 31a Dineura militaris 3Taa H. australis. 36 38(37) 39(37) 40(3) -¥) bb cc op bb cc Marko Prous et al. / Journal of Hymenoptera Research 40: 1-69 (2014) Mesoscutellar appendage 4.0—5.2 times as wide as long. In female, mesepisternum entirely orange; in male, black, or partly or entirely orange. Abdomen often orange. One species of Dineura, D. militaris (Cresson, 1880). Nearctic. sal a Rater Atal oS hae Ane tetas Men Dineura Dahlbom, 1835 in part Mesoscutellar appendage 2.8—3.5 times as wide as long. In female mesepisternum black or partly orange; in male, black. Abdomen black. Some East Palaearctic species of Platycampus: P amurensis Zinovjev, 1986, P coryli Zinovjev, 1986, P japonicus Togashi, 1991. Key to most of the species by Zinovjev (1986). Pera carers Fen leas erecta eres Platycampus Schiodte, 1839 in part Clypeus deeply emarginate. Malar space equal to or slightly longer than diameter of front ocellus. Vein 2r-rs of fore wing present. Twelve species. Holarctic and Oriental. Bie eR Rial tabess PAAR Re Mkt AR ReaD ines una Hemichroa Stephens, 1835 Clypeus shallowly emarginate. Malar space more than 1.5 times as long as diameter of front ocellus. Vein 2r-rs of fore wing absent. Five species of Platycampus. Key to most of the species by Zinovjev (1986). Palaearctic and Oriental. sass hear pabenash td anhe Massa’ Beas Pease Teale Platycampus Schiodte, 1839 in part Notauli weakly outlined Few Pristiphora species. Formerly Pristola. Nearctic. hi Be Beers See ithe RSM clout woes «cae Pristiphora Latreille, 1810 in part Noth Sharp yOrillaned Si... sues seh scue seth oranda este eld sae Sa atau 41 The genera of Nematinae (Hymenoptera, Tenthredinidae) oF 39aa/39bb — 400 Geer eee A0aa eee x ut a = 38a,38b Dineura militaris 38aa,38bb Platycampus amurensis 39al39b,39c Hemichroa australis 39aal39bb,39cc Platycampus luridiventris 40a Pristiphora macnabi 40aa Pseudodineura fuscula. 38 41(40) 42(41) 43(2) 44(43) op op bb cc bb bb Marko Prous et al. / Journal of Hymenoptera Research 40; 1-69 (2014) Malar space shorter than half a diameter of front ocellus. | asl eb fe) ae: de We oe re, ett Re a A EP, Nr Pb 42 Malar space as long as or longer than diameter of front ocellus. Nearctic. Three species. Key to species by Smith (1976b). saith oa he Male ate aacccte het atts ae eee ata ant garctasad th Rina oe cite ete at coatatles Kerita Ross, 1937 Base of vein 2A+3A incomplete and straight. Vein 2r-m usually present. Vein 2m-cu present and joined to vein M either proximal of 2r-m or very slightly distal of 2r-m. Twelve species. Holarctic. pes A ihe Serene te, erred Wreeew ts rere Re Pseudodineura Konow, 1885 Base of vein 2A+3A more or less complete and curved up to 1A. Vein 2r-m of fore wing often absent. Vein 2m-cu absent or present and joined to vein M distal of 2r-m (if 2r-m present; e.g. Fig 43aa). One species, E. anemones (Hering, 1924). West Palaearctic. pene Sucbe te ternal WiRe aed ten ooo ee Endophytus Hering, 1934 in part Base of vein 2A+3A of fore wing abruptly curved up to 1A. Vein 2A of hind wing incomplete, thus cell open. One species, E. anemones (Hering, 1924). West Palaearctic. soa ee ete tte osha a de taedeysniea et Ge Endophytus Hering, 1934 in part Base of vein 2A+3A of fore wing gradually fuses with 1A. Vein 2A of hind wing complete (rarely incomplete in Monocellicampa), TRL sSyer | Mi ber sia [Pao Beet peti I orien Oe, Sie en, ee ade See 44 Length of vein R between junctions with veins M and Rs+M longer than first sector of Rs (see also Fig. 33a). Male and female flagellum similar: thread or seta-like, with short PUD CSEIGES Fras Senn RSet sseevenatervsansahan smeuir vans seartecbeneneae Te 45 Length of vein R between junctions with veins M and Rs+M not longer than first sector of Rs (see also Fig. 33aa). Male and female flagellum dissimilar: seta-like, with short pubescence in female, and seta-, comb-, or saw-like, with long pubescence in male. 41 species. Holarctic and Oriental, introduced to Neotropic and Australasia. seta Seaside terete tla ented saetlandlan deta Cladius \lliger, 1807 in part The genera of Nematinae (Hymenoptera, Tenthredinidae) 39 44bb Ala Pseudodineura fuscula 4\aa Kerita fidala 42al42b/42c P enslini 42aal42bb/42cc (43a) En- dophytus anemones 43aa (44a) Hoplocampa chrysorrhoea 43aa (44aa) Cladius compressicornis 43b E. anemones 43bb Z. chrysorrhoea 44b Platycampus luridiventris 3S, & (left); H. chrysorrhoea 3,8 44bb Cladius grandis 3, 2 (top); C. compressicornis 3, 2 (bottom); C. pectinicornis 3, 2 (right). 40 45(44) 46(45) 47 (46) 48(45) -¥) Oo bb bb Marko Prous et al. / Journal of Hymenoptera Research 40: 1-69 (2014) Vei2 iets Ol Lore WAN DLSGEII cour e wh alll vin cee cxnedep os telaliey telocsgavoh euedaps 46 Nei? Ets tOE OPE WANG ADSEIE tooth cesasayuleonme eat anesd ia cundrnnteaeeeeeutas 48 Combined length of scape and pedicel 0.7 times as long as or longer than flagellomere 1. Epicnemial surface sharply or weakly outlined by furrow................. 47 Combined length of scape and pedicel 0.5 times as long as or shorter than flagellomere 1. Epicnemial surface not outlined. Ten species. Key to species by Smith (2006). Western Nearctic. he tapas ee ae acted as eee nee Susana Rohwer and Middleton, 1932 Vein m-cu of hind wing present, thus cell M closed. Epicnemial surface sharply outlined by furrow. 42 species. Holarctic and Oriental. Re ViaicerR GAA DAW NET? LE ULAR EAR Ce US AURA TUN Hoplocampa Hartig, 1837 Vein m-cu of hind wing absent, thus cell M open. Epicnemial surface weakly outlined by furrow. One species, M. pruni Wei, 1998. East Palaearctic. ons Prec ts te Rh lc A nc afc at ae Monocellicampa Wei, 1998 Combined length of scape and pedicel 0.7 times as long as or longer than flagellomere 1. Pedicel longer than wide. Clypeus with shallow emargination. do cayeaved Fela cqn abel noi) els nag 001 10 aR RNR a en RT ATOR Tee tay 49 Combined length of scape and pedicel usually 0.5 times as long as or shorter than flagellomere 1. Pedicel shorter than wide. Clypeus with deep or shallow emargination. Body: bern oth, APL tid os ceancatatenees state ahead dearest vohaniaMaaudvahenlesonavaait 50 The genera of Nematinae (Hymenoptera, Tenthredinidae) 4] € 46b/47b 46b/47bb —> ¥ st we ‘ 2 = cant - bb? 2 Gea Maer a 45a Hoplocampa chrysorrhoea 45aa Caulocampus acericaulis 46a H. chrysorrhoea 46aa Susana cupressi 46b/47b ZH. chrysorrhoea 46b/47bb Monocellicampa pruni 46bb S. cupressi 41a H. chrysorrhoea 4Taa M. pruni 48al48b C. acericaulis 48aal48bb Cladius compressicornis 48cl48cc C. acericaulis 48cc Cladius compressicornis (left), Anhoplocampa yunanensis (Fig. modified from Wei and Niu 2011). 42 49(48) a bb 50(48) a — aa Marko Prous et al. / Journal of Hymenoptera Research 40; 1-69 (2014) Cell A of hind wing rectangular, widening at apex with vein 1A beginning from anterior margin of cell A. Subapical tooth of tarsal claw long and slender. Two species. Key to species by D.R. Smith (1968). Nearctic. Aer aT eel | 7 dA ma eB Uc ie PER Caulocampus Rohwer, 1912 Cell A of hind wing not rectangular, tapering at apex with vein 1A beginning approximately from centre of cell A. Subapical tooth of tarsal claw absent. One species, A. necopinus (Zhelochovtsev, 1941). West Palaearctic. se ghinte Ach Arbase ease Rashes Aah eer ues Armenocampus Zinovjev, 2000 Epicnemium broad and not elevated, with narrow furrow between the epicnemium and mesepisternum, not strongly curved in upper part of mesepisternum. Sawsheath not distinctly emarginate in dorsal view. Few specimens of Cladius. FW ela Rhea EEN OR AOL UR. Cladius \lliger, 1807 in part Epicnemium narrow and strongly elevated, with broad and deep furrow between epicnemium and mesepisternum, strongly curved in upper part of mesepisternum. bb Sawsheath distinctly emarginate in dorsal view. Three species. Key to species by Wei and Niu (2011). East Palaearctic and Oriental. Be oie Daten oa Meet Ate Uae ara alae a Na Anhoplocampa Wei, 1998 The genera of Nematinae (Hymenoptera, Tenthredinidae) 43 49a,49b Caulocampus acericaulis 49aa Hoplocampa chrysorrhoea 49bb Anoplonyx ovatus 50a,50b Cladius compressicornis 50aa,bb Anhoplocampa yunanensis (Figs modified from Wei and Niu 2011). 44 Marko Prous et al. / Journal of Hymenoptera Research 40: 1-69 (2014) Taxonomy Adelomos Ross, 1935. Neatctic. Adelomos Ross, 1935: 201-202. Type species: Adelomos cleone Ross, 1935, by original designation. Anhoplocampa Wei, 1998. East Palaearctic, Oriental. Anhoplocampa Wei, 1998: 14-15. Type species: Anhoplocampa fumosa Wei, 1998, by original designation. Anoplonyx Marlatt, 1896. Holarctic. Anoplonyx Marlatt, 1896: 18. Type species: Nematus lariciphagus Zaddach in Brischke, 1883 [= Nematus pectoralis auct. nec Lepeletier], by subsequent designation of Taeger et al. (2010: 11). Marlattia Ashmead, 1898: 287. Type species: Hemichroa laricis Marlatt, 1896 [= An- oplonyx luteipes (Cresson, 1880)], by original designation. Armenocampus Zinovjev, 2000. West Palaearctic. Armenocampus Zinovjev, 2000: 452. Type species: Caulocampus necopinus Zhelochovtsev, 1941 [= Armenocampus necopinus (Zhelochovtsev, 1941)], by original designation. Caulocampus Rohwer, 1912. Nearctic. Caulocampus Rohwer, 1912: 239-240. Type species: Priophorus acericaulis MacGillivray, 1906 [= Caulocampus acericaulis (MacGillivray, 1906)], by original designation. Cladius \\liger, 1807. Palaearctic, Nearctic, Oriental, [Neotropical], [Australasian], Fossil. Cladius Mliger, 1807: 190. Type species: Tenthredo difformis Panzer, 1799 [= Cladius pectinicornis (Geoffroy, 1785)], by subsequent designation of Latreille (1810). Nematus (Priophorus) Dahlbom, 1835: 4, 7. Type species: Tenthredo compressicornis Fabricius, 1804 [= Cladius compressicornis (Fabricius, 1804)], by subsequent desig- nation of Blank et al. (2009). Cladius (Trichiocampus) Hartig, 1837: 176. Type species: Nematus grandis Lepeletier, 1823 [= Cladius grandis (Serville, 1823)], by subsequent designation of Rohwer (1911). Stevenia Brullé, 1846: 667. Type species: Pristiphora varipes Lepeletier, 1823 [= Pristiphora varipes Serville, 1823], by monotypy. Note. Correct placement of the name uncertain. ‘The type species “Lep. no. 178” (= Pristiphora varipes Lepeletier) is unplaced. Eudryas Gistel, 1848: viii. Name for Cladius Illiger, 1807. Homonym of Eudryas Boisduval, 1836 [Lepidoptera]. Prionophorus Agassiz, 1848: 888, 889. Name for Nematus (Priophorus) Dahlbom, 1835. Craterocercus Rohwer, 1911. Nearctic. Craterocercus Rohwer, 1911: 385. Type species: Hemichroa phytophagica Dyar, 1898 [= Craterocercus obtusus (Klug, 1816)], by original designation. The genera of Nematinae (Hymenoptera, Tenthredinidae) 45 Dinematus Lacourt, 2006. West Palaearctic. Dinematus Lacoutt, 2006: 237-239. Type species: Dinematus krausi Lacourt, 2006, by original designation. Note. Possibly Pristiphora. Dineura Dahlbom, 1835. Holarctic, Fossil. Tenthredo (Dineura) Dahlbom, 1835: 5, 13. Type species: Tenthredo De Geeri Klug, 1817 [= Dineura virididorsata (Retzius, 1783)], by subsequent designation of Westwood (1839). Leachia Brullé, 1846: 664. Note. Published as a synonym of Dineura Dahlbom as “Leachia Lep. (Mss.)” and never used as valid. Dinevra Agassiz, 1848: 358. Name for Tenthredo (Dineura) Dahlbom, 1835. Hemichroa (Varna) Ross, 1937: 79; syn. n. Type species: Nematus militaris Cresson, 1880 [= Dineura militaris (Cresson, 1880), comb. n.], by original designation. Driocampus J. Zhang & X. Zhang, 1990. Fossil. Driocampus J. Zhang & X. Zhang, 1990: 29-30. Type species: Driocampus shanwan- ganus J. Zhang & X. Zhang, 1990, by original designation. Endophytus Hering, 1934. West Palaearctic. Endophytus Hering, 1934: 353. Type species: Endophytus anemones (Hering, 1924), by original designation. Neopelmatopus Conde, 1934: 181. Type species: Pelmatopus anemones Hering, 1924 [= Endophytus anemones (Hering, 1924)], by original designation. Verna Kaisila, 1960: 300. Type species: Pelmatopus tenuiserra Lindqvist, 1949 [= En- dophytus anemones (Hering, 1924)], by monotypy. Note. Published in synonymy of Pelmatopus. Verna Kontuniemi in Kaisila, 1960: 170-171. Type species: Pelmatopus tenuiserra Lindqvist, 1949 [= Endophytus anemones (Hering, 1924)], by original designation. Eohemichroa Zhelochovtsev & Rasnitsyn, 1972. Fossil. Eohemichroa Zhelochovtsev & Rasnitsyn, 1972: 323. Type species: Hemichroa eophila Cockerell, 1906 [= Eohemichroa eophila (Cockerell, 1906)], by original designation. Euura Newman, 1837. Holarctic, Oriental, [Neotropical], [Afrotropical], [Australasian], Fossil. Euura Newman, 1837: 259-260. Type species: Euura gallae Newman, 1837 [= Euura mucronata (Hartig, 1837)], by subsequent designation of Rohwer (1911). Note. Identity of E. gallae Newman, 1837 is discussed in Liston and Prous (2014). Nematus (Cryptocampus) Hartig, 1837: 221-222. Type species: Nematus (Cryptocampus) medullaris Hartig, 1837 [= Euura amerinae (Linné, 1758)], by subsequent designation of Rohwer (1911). Note. See Opinion 1963 (ICZN 2000). Evura Agassiz, 1848: 439, 440. Name for Euura Newman, 1837. Pontania Costa, 1852. Original paper not seen, cited in Costa (1854): 293; syn. n. Type species: Pontania gallicola Costa, 1852 [= Euura proxima (Serville, 1823), comb. n.], by monotypy. Note. Separatum published in 1852 (Hagen 1862). 46 Marko Prous et al. / Journal of Hymenoptera Research 40: 1-69 (2014) Epitactus Forster, 1854: 435; syn. n. Type species: Epitactus praecox Forster, 1854 [= Euura clitellata (Serville, 1823), comb. n.], by monotypy. Note. Opinion 2280 (ICZN 2011) gives precedence of the name Pachynematus Konow, 1890 over Epitactus, whenever they are consider to be synonyms. Amauronematus Konow, 1890: 233, 237-238, syn. n. Type species: Nematus stenogaster Forster, 1854 [= Euura stenogaster (Forster, 1854), comb. n.], by subsequent des- ignation of Lacourt (1999). Holcocneme Konow, 1890: 233, 238, syn. n. Type species: Nematus vicinus Serville, 1823 [= Euura vicina (Serville, 1823), comb. n.], by subsequent designation of Blank et al. (2009). Pachynematus Konow, 1890: 233, 238, syn. n. Type species: Nematus trisignatus Forster, 1854 [= Euura clitellata (Serville, 1823), comb. n.], by subsequent designation of Schmidt et al. (1998). Brachycolus Konow, 1895: 166, syn. n. Homonym of Brachycolus Buckton, 1879 [He- miptera]. Type species: Nematus viduatus (Zetterstedt, 1838) [= Euura viduata (Zetterstedt, 1838), comb. n.], by subsequent designation of Rohwer (1911b). Nematus (Holcocnema) Schulz, 1906: 80, syn. n. Name for Holcocneme Konow, 1890. Holcocnemis Konow, 1907: 331, syn. n. Name for Holcocneme Konow, 1890. Pteronidea Rohwer, 1911: 98, syn. n. Type species: Nematus ventralis Say, 1824 [= Euura ventralis (Say, 1824), comb. n.], by original designation. Note. Proposed as new name for Pteronus sensu Panzer (1806). Pontopristia Malaise, 1921: 12-13, syn. n. Type species: Nematus suavis Ruthe, 1859 [= Euura amentorum (Forster, 1854), comb. n.], by original designation. Brachycoluma Strand, 1929: 26, syn. n. Name for Brachycolus Konow, 1895. Decanematus Malaise, 1931: 31, syn. n. Type species: Decanematus longiserra Malaise, 1931 [= Euura malaisei (Hellén, 1970), comb. n.], by original designation. Pikonema Ross, 1937: 86, syn. n. Type species: Nematus dimmockii Cresson, 1880 [= Euura dimmockii (Cresson, 1880), comb. n.], by original designation. Phyllocolpa Benson, 1960: 60, syn. n. Type species: Nematus leucapsis Tischbein, 1846 [= Euura leucapsis (Tischbein, 1846), comb. n.], by original designation. Eitelius Kontuniemi, 1966: 44-47, syn. n. Type species: Pachynematus dentatus Lindqvist, 1937 [= Euura dentata (Lindqvist, 1937), comb. n.], by original designation. Euura (Gemmura) E.L. Smith, 1968: 1401. Type species: Cryptocampus mucronatus (Hartig, 1837) [= Euura mucronata (Hartig, 1837)], by original designation. Note. See Opinion 1963 (ICZN 2000). Ribinematus Kontuniemi, 1975: 38, syn. n. Not available. Type species: no type species selected. Pontania (Eupontania) Zinovjev, 1985: 4, syn. n. Type species: Nematus vesicator Bremi- Wolf, 1849 [= Euura vesicator (Bremi-Wolf, 1849), comb. n.], by original designation. Nematus (Larinematus) Zhelochovtsev in Zhelochovtsev and Zinovjev, 1988: 70 (key), 169, syn. n. Type species: Nematus imperfectus Zaddach, 1876 [= Euura imperfecta (Zaddach, 1876), comb. n.], by original designation. Nematus (Polynematus) Zhelochovtsev in Zhelochovtsev and Zinovjev, 1988: 71 (key), 137, syn. n. Type species: applying Article 70.3.2 (ICZN 1999), we hereby select as type The genera of Nematinae (Hymenoptera, Tenthredinidae) 47 species Nematus annulatus Gimmerthal, 1834 [= Euura annulata (Gimmerthal, 1834), comb. n.], this being the taxonomic species indicated by Zhelochovtsev’s original desig- nation of “Nematus rumicis L.” [misidentification of Tenthredo rumicis Linnaeus, 1758; currently treated as an unplaced name in the Tenthredinidae: Taeger et al. 2010]. Nematus (Bacconematus) Zhelochovtsev in Zhelochovtsev & Zinovjev, 1988: 70 (key), 128, syn. n. Type species: Nematus pumilio (Konow, 1903) [= Euura pumilio (Konow, 1903), comb. n.], by original designation. Alpinematus Lacoutrt, 1996: 269-270, syn. n. Type species: Alpinematus elongatulus La- court, 1996 [= Euura elongatula (Lacourt, 1996), comb. n.], by original designation. Epicenematus Lacoutrt, 1998: 82, syn. n. Type species: Nematus pallescens Hartig, 1837 [= Euura pallescens (Hartig, 1837), comb. n.], by original designation. Kontuniemiana Lacoutt, 1998: 80-81, syn. n. Type species: Tenthredo ribesii Scopolii, 1763 [= Euura ribesii (Scopoli, 1763), comb. n.], by original designation. Lindqvistia Lacourt, 1998: 81, syn. n. Type species: Nematus reticulatus Holmgren, 1883 [= Euura reticulata (Holmgren, 1883), comb. n.], by original designation. Luea Wei and Nie, 1998: 15-18, syn. n. Type species: Luea sinica Wei and Nie, 1998 [= Euura sinica (Wei and Nie, 1998), comb. n.], by original designation. Tubpontania Vikberg, 2010: 4, syn. n. Type species: Nematus anomalopterus Forster, 1854 [= Euura anomaloptera (Forster, 1854), comb. n.], by original designation. Fagineura Vikberg & Zinovjev, 2000. East Palaearctic. Fagineura Vikberg & Zinovjev in Shinohara et al., 2000: 114-115. Type species: Fagineura crenativora Vikberg and Zinovjev, 2000, by original designation. Fallocampus Wong, 1977. Nearctic. Fallocampus Wong, 1977: 1103-1107. Type species: Camponiscus americanus Marlatt, 1896 [= Fallocampus americanus (Marlatt, 1896)], by original designation. Florissantinus Zhelochovtsev & Rasnitsyn, 1972. Fossil. Florissantinus Zhelochovtsev and Rasnitsyn, 1972: 320-323. Type species: Florissanti- nus angulatus Zhelochovtsev and Rasnitsyn, 1972, by monotypy. Hemichroa Stephens, 1835. Holarctic, Oriental. Hemichroa Stephens, 1835: 55. Type species: Tenthredo australis Serville, 1823 [= Hemi- chroa australis (Serville, 1823)], by subsequent designation of Blank et al. (2009). Dineura (Leptocerca) Hartig, 1837: 228. Type species: Tenthredo alni Linné, 1767 [= Hemichroa australis (Serville, 1823)], by subsequent designation of Rohwer (1911). Engages Gistel, 1848: ix. Name for Dineura (Leptocerca) Hartig, 1837. Leptocercus Thomson, 1871: 69, 70, 76. Name for Dineura (Leptocerca) Hartig, 1837. Hoplocampa Hartig, 1837. Holarctic, Oriental, Fossil. Tenthredo (Hoplocampa) Hartig, 1837: 276-277. Type species: Tenthredo (Allantus) brevis Klug, 1816 [= Hoplocampa brevis (Klug, 1816)], by subsequent designation of Rohwer (1911). Note. Described as Tenthredo (Selandria |Hoplocampa)). 48 Marko Prous et al. / Journal of Hymenoptera Research 40: I-69 (2014) Macgillivraya Ashmead, 1898: 257. Homonym of Macgillivraya Forbes, 1852 [Mol- lusca]. Type species: Macgillivraya oregonensis Ashmead, 1898 [= Hoplocampa oregonensis (Ashmead, 1898)], by original designation. Macgillivrayella Ashmead, 1900: 606. Name for Macgillivraya Ashmead, 1898. Katsujia Togashi, 1964. East Palaearctic. Katsujia Togashi, 1964: 479. Type species: Katsujia planaritibia Togashi, 1964, by original designation. Kerita Ross, 1937. Nearctic. Kerita Ross, 1937: 80. Type species: Kerita fidala Ross, 1937, by original designation. Megadineura Malaise, 1931. East Palaearctic, Oriental. Megadineura Malaise, 1931: 147-148. Type species: Dineura grandis André, 1882 [= Megadineura grandis (André, 1882)], by original designation. Stenomesoneura Wei, 1998: 411. Not available. Type species: Stenomesoneura apicalis Wei, 1998 [= Megadineura grandis (André, 1882)], by monotypy. Note. Not avail- able since only pictures and no description in words are included in the work. Mesoneura Hartig, 1837. Palaearctic. Dineura (Mesoneura) Hartig, 1837: 228-229. Type species: Tenthredo opaca Fabricius, 1775 [= Mesoneura opaca (Fabricius, 1775)], by subsequent designation of Taeger and Blank (1996). Selandria (Pristis) Brullé, 1846: 665. Type species: Tenthredo punctigera Lepeletier, 1823 [= Mesoneura opaca (Fabricius, 1775)], by original designation. Mesonevra Agassiz, 1848: 667. Name for Dineura (Mesoneura) Hartig, 1837. Lisconeura Rohwer, 1908: 529. Type species: Scolioneura vexabilis Brues, 1908 [= Mesoneura vexabilis (Brues, 1908)], by original designation. Monocellicampa Wei, 1998. East Palaearctic. Monocellicampa Wei, 1998: 16. Type species: Monocellicampa pruni Wei, 1998, by original designation. Moricella Rohwer, 1916. Oriental. Moricella Rohwer, 1916: 111. Type species: Moricella rufonota Rohwer, 1916, by original designation. Nematinus Rohwer, 1911. Holarctic. Nematinus Rohwer, 1911: 84. Type species: Nematus fuscipennis Lepeletier, 1823 [= Nematinus fuscipennis (Serville, 1823)], by subsequent designation of Taeger and Blank (1996). The genera of Nematinae (Hymenoptera, Tenthredinidae) 49 Nematus Panzer, 1801. Holarctic, Oriental. Nematus Panzer, 1801: 82:10. Type species: Tenthredo (Nematus) lucida Panzer, 1801 [= Nematus lucidus (Panzer, 1801)], by monotypy. Note. Described in synonymy of Tenthredo lucida. Nematus Jurine in Panzer, 1801: 163. Type species: no type species selected. Note. Suppressed by Opinion 135 (ICZN 1939). Nematus Jurine, 1807: 59. Homonym of Nematus Panzer, 1801. Type species: no type species selected. Craesus Leach, 1817: 129. Type species: Nematus septentrionalis (Linné, 1758), by monotypy. Hypolaepus WF. Kirby, 1882: 324-325. Type species: Hypolaepus abbotii W.F. Kirby, 1882 [= Nematus abbotii (W.F. Kirby, 1882)], by monotypy. Nematus (Paranematus) Zinovjev, 1978: 626-627. Type species: Nematus wahlbergi Thomson, 1871, by original designation. Neodineura Vaeger, 1989. West Palaearctic. Neodineura Yaeger, 1989: 150-151. Type species: Mesoneura arquata (Klug, 1816) [= Neodineura arquata (Klug, 1816)], by original designation. Nescianeura Lacourt, 2006. West Palaearctic. Paraneura Lacoutrt, 2004: 42. Not available. Nomen nudum. Nescianeura Lacourt, 2006: 235-236. Type species: Nescianeura noblecourti Lacourt, 2006, by original designation. Note. Probably Euura or Nematus. Platycampus Schiédte, 1839. Palaearctic, Oriental. Nematus (Leptopus) Hartig, 1837: 184. Homonym of Leptopus Latreille, 1809 [Hemip- tera]. Type species: Nematus hypogastricus Hartig, 1837 [= Platycampus luridiventris (Fallén, 1808)], by monotypy. Platycampus Schigdte, 1839: 20. Name for Nematus (Leptopus) Hartig, 1837. Erasminus Gistel, 1848: ix. Name for Nematus (Leptopus) Hartig, 1837. Camponiscus Newman, 1869: 215-217. Type species: Camponiscus healaei Newman, 1869 [= Platycampus luridiventris (Fallén, 1808)], by monotypy. Pristiphora Latreille, 1810. Holarctic, Oriental, Neotropical. Pristiphora Latreille, 1810: 294, 435. Type species: Pteronus testaceus Jurine, 1807 [= Pristiphora testacea (Jurine, 1807)], by original designation. Nematus (Diphadnus) Hartig, 1837: 225. Type species: Nematus fuscicornis Hartig, 1837 [= Pristiphora appendiculata (Hartig, 1837)], by subsequent designation of Gimmerthal (1847). Lygaeonematus Konow, 1890: 233, 238. Type species: Nematus pini (Retzius, 1783) [= Pristiphora abietina (Christ, 1791)], by subsequent designation of Rohwer (1911). Micronematus Konow, 1890: 233, 239. Type species: Nematus pullus Forster, 1854 [= Pristiphora monogyniae (Hartig, 1840)], by subsequent designation of Rohwer (1911). 50 Marko Prous et al. / Journal of Hymenoptera Research 40: 1-69 (2014) Gymnonychus Marlatt, 1896: 19 (key), 122. Type species: Gymnonychus californicus Marlatt, 1896 [= Pristiphora abbreviata (Hartig, 1837)], by original designation. Neopareophora MacGillivray, 1908: 289. Type species: Neopareophora martini MacGillivray, 1908 [= Pristiphora litura (Klug, 1816), comb. n.], by original designation. Neotomostethus MacGillivray, 1908: 290. Type species: Neotomostethus hyalinus MacGil- livray, 1908 [= Pristiphora hyalina (MacGillivray, 1908)], by original designation. Dineuridea Rohwer, 1912: 240. Type species: Marlattia erythrothorax Rohwer, 1911 [= Pristiphora erythrothorax (Rohwer, 1911), comb. n.], by original designation. Pristiphora (Sala) Ross, 1937: 85. Type species: Nematus chloreus Norton, 1867 [= Pristiphora chlorea (Norton, 1867)], by original designation. Pristola Ross, 1945: 153-154, syn. n. Type species: Pristola macnabi Ross, 1945 [= Pristiphora macnabi (Ross, 1945), comb. n.], by original designation. Lygaeonematus (Lygaeotus) Lindqvist, 1952: 82. Not available. Type species: no type species selected. Lygaeonematus (Lygaeophora) Lindqvist, 1952: 82. Not available. Type species: no type species selected. Nepionema Benson, 1960: 173-174, syn. n. Type species: Nepionema helvetica Benson, 1960 [= Pristiphora helvetica (Benson, 1960), comb. n.], by original designation. Melastola Wong, 1968: 1049, 1051, 1053, syn. n. Type species: Gymnonychus resinicolor Marlatt, 1896 [=Pristiphora resinicolor (Marlatt, 1896)], by original designation. Sharliphora Wong, 1969: 332-334. Type species: Tenthredo ambiguus Fallén, 1808 [= Pristiphora nigella (Forster, 1854)], by original designation. Lygaeophora Hellén, 1975: 100. Not available. Type species: Nematus lanificus Zaddach in Brischke 1883 [= Pristiphora lanifica (Zaddach, 1883)], by original designation. Note. Treated as a junior synonym of Pristiphora and therefore not made available (Art. 11.5., ICZN 1999). Lygaeotus Hellén, 1975: 100. Not available. Type species: Nematus coactulus Ruthe, 1859 [= Pristiphora coactula (Ruthe, 1859)], by original designation. Note. Invalid because published as a junior synonym of Pristiphora. Pristiphora (Lygaeophora) Liston, 1981: 181-184. Not available. Type species: no type species selected. Nematus (Oligonematus) Zhelochovtsev in Zhelochovtsev and Zinovjev, 1988: 72 (key), 162. Type species: Nematus laricis Hartig, 1837 [= Pristiphora laricis (Hartig, 1837)], by original designation. Pristiphora (Lygaeotus) Liston, 1993: 105. Type species: Nematus coactulus Ruthe, 1859 [= Pristiphora coactula (Ruthe, 1859)], by original designation. Pristiphora (Lygaeophora) Liston, 1993: 104-105. Type species: Lygaeonematus variipes Lindqvist, 1952 [= Pristiphora sermola Liston, 1993], by original designation. Pristicampus Zinovjev, 1993: 80, syn. n. Type species: Mesoneura arctica Lindqvist, 1959 [= Pristiphora arctica (Lindqvist, 1959), comb. n.], by original designation. The genera of Nematinae (Hymenoptera, Tenthredinidae) 54 Pseudodineura Konow, 1885. Holarctic. Dolerus (Pelmatopus) Hartig, 1837: 244. Homonym of Pelmatopus Fischer von Waldheim, 1824 [Coleoptera]. Type species: Dolerus minutus Hartig, 1837 [= Pseudodineura fuscula (Klug, 1816)], by monotypy. Pseudodineura Konow, 1885: 295, 297. Type species: Tenthredo (Allantus) parvula Klug, 1816 [= Pseudodineura parvula (Klug, 1816)], by subsequent designation of Rohwer (1911). Phyllopais Hering, 1934: 353. Name for Dolerus (Pelmatopus) Hartig, 1837. Renonerva Wei & Nie, 1998. East Palaearctic. Renonerva Wei & Nie, 1998: 14-15. Type species: Renonerva fumosa Wei and Nie, 1998, by original designation. Stauronematus Benson, 1953. Palaearctic. Stauronema Benson, 1948: 22. Homonym of Stauronema Sollas, 1877 [Spongidae]. Type species: Nematus platycerus Hartig, 1840 [= Stauronematus platycerus (Hartig, 1840)], by subsequent designation of Liston (2007). Stauronematus Benson, 1953: 153. Name for Stauronema Benson, 1948. Susana Rohwer & Middleton, 1932. Nearctic. Susana Rohwer & Middleton, 1932: 93. Type species: Susana cupressi Rohwer and Middleton, 1932, by original designation. Unplaced Nematinae genera Messa Leach, 1817 West Palaearctic Messa Leach, 1817: 126. Type species: Messa hortulana Leach, 1817, by monotypy. Secondary homonymy of species names The new and much wider circumscription of Euura adopted in this work involves the synonymy of several partly species-rich nominal genera. As a result, a number of species names become secondary homonyms when they are placed in Euura. In all except five of the 25 cases listed below, the senior homonym has been applied after 1899 to a taxon considered to be valid. Article 23.9.1 of the International Code of Zoological Nomen- clature (ICZN 1999) is therefore not fulfilled in these cases, and the junior homonyms require replacement. In the five remaining cases (Amauronematus poppii, Euura cinereae, Pontopristia punctulata, Pteronidea brachycera and P curticornis) the junior homonym has not been used for a particular taxon, as its presumed valid name, in at least 25 works, published by at least 10 authors in the immediately preceding 50 years and encompass- ing a span of not less than 10 years. The second condition stipulated by Article 23.9.1 is 52 Marko Prous et al. / Journal of Hymenoptera Research 40; 1-69 (2014) thus not met, and these homonyms also require replacement. Only dealt with below are cases of secondary homonymy where the taxonomy of both species at present seems rea- sonably clear. In several remaining cases, the validity of the species denoted by the junior homonym is highly questionable and it is therefore not desirable to replace them now. The decision on whether a replacement name is necessary should be made once the rel- evant groups have been better studied. The replacement names proposed are suggested by the authors of the present paper who studied the individual cases, except for the species described in Euura, Phyllocolpa and Pontania. Jens-Peter Kopelke allowed us to publish, in his own words, the relevant replacement names for secondary homonyms of the latter. Amauronematus acutiserra Lindqvist, 1974; secondary homonym of Pontania acutiserra Lindqvist, 1949, currently recognised as valid in the combination Euura acutiserra (Lindqvist, 1949), comb. n. Euura aceroserra Taeger and Blank, nom. n. is pro- posed for A. acutiserra Lindqvist. Amauronematus atratus Lindqvist, 1961; secondary homonym of Pontania atrata MacGillivray, 1919, currently recognised as valid in the combination Euura atrata (MacGillivray, 1919), comb. n. Euura lethe Prous and Liston, nom. n. is proposed for A. atratus Lindqvist. Amauronematus enslini Lindqvist, 1959; secondary homonym of Pontania enslini Zirngiebl, 1937, currently recognised as a junior subjective synonym of Euura crassipes (Thomson, 1871), comb. n. Euura bavarica Blank and Liston, nom. n. is proposed for A. enslini. Amauronematus lateralis Konow, 1895; secondary homonym of Nematus lateralis Norton, 1867, currently recognised as valid in the combination Euura lateralis (Norton, 1867), comb. n. The valid name for A. /ateralis Konow is E. trautmanni (Enslin, 1919). Amauronematus mimus Schmidt, 1997; secondary homonym of Preronus mimus Konow, 1903, currently recognised as valid in the combination Euura mima (Konow, 1903), comb. n. Euura mimator Schmidt, nom. n. is proposed for A. mimus Schmidt. Amauronematus nitens Lindqvist, 1977; secondary homonym of Nematus nitens Thom- son, 1888, currently recognised as a junior subjective synonym of Euura respondens (Forster, 1854), comb. n. Euura nitidula Prous and Taeger, nom. n. is proposed for A. nitens Lindqvist. Amauronematus pacificus Malaise, 1931; secondary homonym of Pontania pacifica Marlatt, 1896, currently recognised as valid in the combination Euura pacifica (Marlatt, 1896), comb. n. Amauronematus obscurus Lindqvist, 1962 is a junior subjective synonym of E. pacifica (Malaise) but the former is not available as a replacement name because it is a junior secondary homonym of Euura obscura (Norton, 1861), comb. n. Euura tranquilla Vardal and Prous, nom. n. is proposed for A. pacificus Malaise. Amauronematus poppii Konow, [September]1904; secondary homonym of Pontania poppii Konow, [July] 1904, currently recognised as a junior subjective synonym of Euura parvula (Holmgren, 1883), comb. n. Euura bertilpoppii Heibo and Liston, nom. n. is proposed for A. poppii Konow. The genera of Nematinae (Hymenoptera, Tenthredinidae) 53 Amauronematus propinquus Saarinen, 1950; secondary homonym of Cryptocampus pro- pinquus Rohwer, 1909, currently recognised as valid in the combination Euura propinqua (Rohwer, 1909), comb. n. Euura propinquator Schmidt, nom. n. is proposed for A. propinquus Saarinen. Euura (Gemmura) boreoalpina Kopelke, 2001; secondary homonym of Amauronema- tus (Pontopristia) boreoalpina (Lindqvist, 1961), currently recognised as valid in the combination Euura boreoalpina (Lindqvist, 1961), comb. n. Euura glaucatumida Kopelke, nom. n. is proposed for FE. boreoalpina Kopelke. Euura (Euura) lapponica Kopelke, 1996; secondary homonym of Pontania (Euponta- nia) lapponica Malaise, 1921, currently recognised as a junior subjective synonym of Euura crassipes (Thomson, 1871). Euura salicislapponicae Kopelke, nom. n. is proposed for E. lapponica Kopelke. Euura (Gemmura) phylicifoliae Kopelke, 2001; secondary homonym of Pontania phyl- icifoliae Forsius, 1919, currently recognised as a junior subjective synonym of Eu- ura arcticornis (Konow, 1904), comb. n. Euura salicisphylicifoliae Kopelke, nom. n. is proposed for E. phylicifoliae Kopelke. Euura (Euura) purpureae Kopelke, 1996; secondary homonym of Nematus purpureae Cameron, 1884, currently recognised as valid in the combination Euura purpureae (Cameron, 1884), comb. n. Euura salicispurpureae Kopelke, nom. n. is proposed for E. purpureae Kopelke. Pachynematus tenuiserra Lindqvist, 1949; secondary homonym of Amauronematus ten- uiserra Lindqvist, 1944, currently recognised as valid in the combination Euura tenuiserra (Lindqvist, 1944), comb. n. Euura scandica Vardal and Heibo, nom. n. is proposed for P. tenuiserra Lindqvist. Phyllocolpa pschornwalcheri Kopelke, 2007; secondary homonym of Nematus pschorn- walcheri Muche, 1972, currently recognised as a junior subjective synonym of Eu- ura monticola (Thomson, 1871), comb. n. Euura hubertpschornwalcheri Kopelke, nom. n. is proposed for P. pschornwalcheri Kopelke. Pontania hastatae Vikberg, 1970; secondary homonym of Euura hastatae Malaise, 1921, currently recognised as valid. Euura hastatavora Vikberg, nom. n. is pro- posed for Pontania hastatae Vikberg. Pontania (Eupontania) montivaga Kopelke, 1991; secondary homonym of Pachynematus montivagus Marlatt, 1896, currently recognised as valid in the combination Euura montivaga (Marlatt, 1896), comb. n. Euura foetidatumida Kopelke, nom. n. is pro- posed for P. montivaga Kopelke. Pontania obscura Kopelke, 2005; secondary homonym of Nematus obscurus Norton, 1861, currently recognised as valid in the combination Euura obscura (Norton, 1861). Euura abdita Kopelke, nom. n. is proposed for P. obscura Kopelke. Pontopristia montana Lindqvist, 1961; secondary homonym of Nematus montanus Zaddach, 1883, currently recognised as valid in the combination Euura montana (Zaddach, 1883), comb. n. Euura oreophila Liston and Prous, nom. n. is proposed for P. montana Lindqvist. 54 Marko Prous et al. / Journal of Hymenoptera Research 40: I-69 (2014) Pontopristia punctulata Lindqvist, 1961; secondary homonym of Nematus punctulatus Thomson, 1863, currently recognised as a junior subjective synonym of Euura vaga (Fabricius, 1781), comb. n. Euura suecica Blank and Taeger, nom. n. is proposed for P. punctulata Lindqvist. Pteronidea brachycera Lindqvist, 1975; secondary homonym of Nematus brachycerus Hartig, 1840, currently recognised as a junior subjective synonym of Euura fallax (Serville, 1823), comb. n. Euura brevicera Taeger and Blank, nom. n. is proposed for P. brachycera Lindqvist. Pteronidea brunnea Lindqvist, 1971; secondary homonym of Nematus brunneus Norton, 1864, currently recognised as valid in the combination Euura brunnea (Norton, 1864), comb. n. The valid name for P. brunnea Lindqvist is E. brunnescens (Vikberg, 1982), comb. n. Pteronidea curticornis Lindqvist, 1969; secondary homonym of Nematus curticornis Cameron, 1885, currently recognised as a junior subjective synonym of Euura pedunculi (Hartig, 1837), comb. n. The valid name for P. curticornis Lindqvist is E. truncicornis (Vikberg, 1982), comb. n. Pteronidea polita Lindqvist, 1974; secondary homonym of Nematus politus Zaddach, 1883, currently recognised as valid in the combination Euura polita (Zaddach, 1883), comb. n. The valid name for P. polita Lindqvist is E. glabra (Vikberg, 1982), comb. n. Discussion It has been evident since 2006 (Nyman et al. 2006) that changes are necessary in the classification of the Nematinae to reflect the current understanding of their phylogeny. Most problematic are the ‘higher’ Nematinae as defined in Nyman et al. (2006). Sub- sequent analyses (Nyman et al. 2010; this work), including more taxa and genes, have confirmed the existence of two well-supported and species-rich clades within the “high- er’ Nematinae. Although there are several ways to divide the phylogenetic tree into gen- era, we decided to treat these two major clades as Pristiphora and Euura, which are the oldest available genus group names for these clades. Although it might seem preferable to use the name Nematus instead of Euura, because most of the species have been placed in Nematus in the past (e.g. Zhelochovtsev and Zinovjev 1988), the phylogenetic place- ment of the type species Nematus lucidus is unfortunately not stable. Previous analyses (Nyman et al. 2006; 2010) placed NV. lucidus closer to Euura, but this relationship is no longer supported in our analysis that also includes NaK sequences. Even if later analyses with more markers support a NV. /ucidus-Euura clade, and Euura could be subsumed under Nematus, the possibility still remains to treat these potential sister groups (it is unlikely that NV. ucidus falls within Euura) as separate genera (Fig. 6). Regardless of the position of NV. lucidus, Euura is a well-supported clade and therefore is a good candidate for a genus. The deeper relationships within this clade, however, are genetically poorly resolved and often at odds with current nomenclature, which argues against creating or The genera of Nematinae (Hymenoptera, Tenthredinidae) 55 Nematus s.1. Nematus Nematus s.s. Euura Pristiphora Euura Pristiphora Euura Pristiphora Nematus this study Nyman et al. 2006; 2010 hypothetical Figure 6. Three possible relationships between the Euura, Pristiphora, and Nematus clades. All are con- sistent with treatment of these three clades as separate genera, or combining all of them as a single genus Nematus. Including Euura, but not Pristiphora, within Nematus is consistent only with the tree shown in the middle (supported by previous analyses: Nyman et al. 2006; 2010). maintaining genera within this clade. Within the Euura clade, another possibility would be to retain monophyletic genera, modify those which are not monophyletic, and create new ones for the remaining species. However, this option would result in the prolif- eration of morphologically and genetically poorly defined, unstable genera (many of which would be monotypic) because of weakly established phylogenetic relationships. Attempting to maintain the status quo is equally unlikely to result in nomenclatural sta- bility, because the creation of new genera and the redefinition of established ones would be expected to continue in the foreseeable future. It would also be possible to include Pristiphora, Mesoneura, Fagineura, and Euura in Nematus (this clade is well supported), but this would result in many more nomenclatural changes (new combinations and secondary homonyms) compared to the “Euura” option. In the other big clade, Pristiphora, which is here treated as a genus, the nomenclatu- ral changes are fortunately less extensive than in Euura. Splitting up Pristiphora in order to maintain a few small genera, such as Melastola, Pristola, Nepionema, Neopareophora and Pristicampus, would cause problems similar to those that would result from splitting up Euura. The creation of several poorly defined genera with weakly resolved phyloge- netic relationships would certainly lead to nomenclatural instability in the future. We want to emphasize that although there are uncertainties regarding many rela- tionships in the phylogeny of Nematinae (including the exact placement of N. lucidus), this does not mean that all or even most relationships are unreliably reconstructed. The support for the two largest clades, Pristiphora and Euura, has increased with the addi- tion of NaK (compared to Nyman et al. 2010, ML bootstrap support went up from 98% to 100% for Euura and from 89% to 99% for Pristiphora). Some taxa in Euura, Nematus, and Pristiphora are monophyletic and relatively well defined (Euura s.s., a large part of Amauronematus, Craesus, Pristicampus etc.), which would allow recognition of these groups. However, for the sake of nomenclatural sta- bility we strongly recommend that the formal acceptance of these clades as genera should be avoided. Preferably such infrageneric groups should be referred to as species groups (within the context of ICZN) or defined according to the PhyloCode (Cantino and de Queiroz 2010). 56 Marko Prous et al. / Journal of Hymenoptera Research 40; 1-69 (2014) Although the Nematus clade received less support than Euura and Pristiphora, we did not attempt to divide it further because it includes only few species and most of them already have the generic name Nematus. The only exception is Craesus as it is usu- ally considered to represent a separate genus. However, it is nearly indistinguishable from the erythrogaster-group of Nematus when we exclude the exceptionally expanded hind tibia and metatarsomere 1 of Craesus (Smith 2008). If future phylogenetic re- search shows that this group is not monophyletic, Nematus can, if necessary, still be divided into smaller genera. However, this will affect only a small number of species. The rather broad definitions of genera like Pristiphora and Euura, and the resulting nomenclatural changes will certainly cause controversy, but in our opinion this is the best solution in light of current evidence. Pristiphora and Euura are strongly supported clades (and it is unlikely that future data will challenge this), but their relationships to other ‘higher’ Nematinae, as well as basal branching patterns within Pristiphora and Euura, remain controversial. We wish to stabilize the nomenclature of ‘higher’ Ne- matinae by considering these clades as genera, so that in the future it is possible to con- centrate on actually studying these sawflies without having to deal with constant name changes. The downside of our approach is that we have to make many nomenclatural changes, especially concerning Euura, but when accepted, it will be a once-only event. The other change, compared to the World Catalog (Taeger et al. 2010), is the transfer of Nematus militaris Cresson, 1880 to Dineura, creating Dineura militaris (Cresson, 1880), comb. n. (previously in Hemichroa). According to DNA sequence data, Dineura militaris forms a clade with Dineura s.s. to the exclusion of other genera in Dineurini (Figs 2 and 5). Absence of a velum in Dineura militaris and other Dineura might be a synapomorphy for these taxa. Based on current data, the higher level relationships (i.e. between genera) within Nematinae are generally not well supported, but there are few clades worth mention- ing: Dineurini, Pseudodineurini and ‘higher’ Nematinae (Fig. 2). Of these three clades, composition of only Pseudodineurini matches with the morphology based classifications (e.g. Zhelochovtsev and Zinovjev 1988; Goulet 1992). Composition of Dineurini and ‘higher’ Nematinae is the same as found by Nyman et al. (2006). However, before any tribal classification of Nematinae is proposed, molecularly yet unsampled genera and more sequence data should be gathered to confidently resolve the phylogeny of genera. Although we are far from having resolved all problems involved in the definition and identification of the genera of Nematinae, we are confident that our pictorial approach using photographs of actual specimens rather than drawings makes identification much easier than before in this challenging group. We regard the key as a starting point for future improvements: the main remaining shortcomings involve Euura, Nematus and Pristiphora, two of which (Euura and Pristiphora) comprise over 75% of nematine spe- cies. Nevertheless, most species of these genera should key out correctly, as only a few Pristiphora species are likely to run to Euura or vice versa. Pristiphora has campaniform sensilla on the tangium of the lancet and generally a more or less truncate clypeus and a swollen apex of vein C, while Euura and Nematus lack campaniform sensilla and gener- ally have an emarginate clypeus and a less swollen apex of vein C. While some groups of The genera of Nematinae (Hymenoptera, Tenthredinidae) 57 Nematus are differentiated morphologically from Euura by characters of metatarsomere 1 (see couplets 12—13 in the Key), we are currently unaware of unequivocal characters to distinguish the Nematus wahlbergi and erythrogaster groups from Euura. In addi- tion, there are currently many, mostly Nearctic species that are placed in Nematus or Pteronidea in Yaeger et al. (2010), but which cannot be associated confidently with our current concepts of Nematus or Euura due to lack of recent revisions and/or DNA sequence data. Solving these problems requires species-level revisions, regardless of how genera are defined. For Euura, Nematus and Pristiphora, a key to species level is urgently needed, rather than having an intermediate step keying out genera or subgenera. Work is currently under way in this direction for the West Palaearctic species. Acknowledgements Funding by the Swedish Taxonomy Initiative (contract number dha 153/2011) made this work possible. We offer our heartfelt thanks for loans and gifts of material, as well as for valuable advice and information, to liro Kakko, Jens-Peter Kopelke, Manfred Kraus, Jean Lacourt, Pekka Malinen, Gengyun Niu, Henri Savina, Akihiko Shinohara, Shu-Jun Wei and staff of the Swedish Malaise Trap Project (particularly Mattias For- shage, Kajsa Glemhorn, Dave Karlsson and Pelle Magnusson). Julie Stahlhut, Crystal Sobel and their colleagues at BOLD (The Barcode of Life Data System, University of Guelph, Canada) helped us to obtain barcode sequences of some specimens. Gavin Broad and Lars Vilhelmsen are cordially thanked for reviewing the manuscript. References Agassiz JLR (1848) Nomenclatoris zoologici index universalis, continens nomina systematica classium, ordinum, familiarum et generum animalium omnium, tam viventium quam fos- silium, secundum ordinem alphabeticum unicum disposita, adjectis homonymiis plan- tarum. Soloduri, Jent et Grassmann, 1135 pp. André E (1882) Notes hyménoptérologiques. 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