Corresponding author: Marko Prous (
Academic editor: Stephan M. Blank
Keys to adults and larvae of the genera of West Palaearctic nematine sawflies are presented. Species of some of the smaller genera are keyed, and their taxonomy, distribution, and host plants reviewed, with a geographic focus on north-western Europe, particularly Sweden.
Prous M, Liston A, Kramp K, Savina H, Vårdal H, Taeger A (2019) The West Palaearctic genera of Nematinae (Hymenoptera, Tenthredinidae). ZooKeys 875: 63–127.
In 2012 a project funded by the Swedish Taxonomy Initiative was launched, with the main objective of improving our knowledge of the taxonomy and distribution of nematine sawflies in Fennoscandia, and Sweden in particular (STI
The Swedish Malaise Trap Project is abbreviated to
In the specimen data the dates are given as dd.mm.yyyy, and coordinates as positive (north or east) or negative (south or west) decimal degrees latitude and longitude.
Morphological terminology mostly follows
First drafts of the key to larvae were based mainly on
DNA was extracted and purified with an EZNA Tissue DNA Kit (Omega Bio-tek) according to the manufacturer’s protocol and stored at -20 °C for later use. Typically, one or two legs were used for DNA extraction, but for males the whole genital capsule was often additionally used to increase DNA yield and to free penis valves from muscles before photography. In some cases, the whole specimen was used for extraction. One mitochondrial and four nuclear regions were used in the phylogenetic analyses, although not all of these genes were obtained for all species. Primers used for amplification and sequencing are listed in Table
Primers used for PCR and sequencing (preferred primers in bold), with information provided on respective gene fragment, primer name, direction (forward, F or reverse, R), primer sequence, standard PCR annealing temperature, utilization (PCR/ sequencing), and reference. Primer annealing temperatures used for sequencing at Macrogen were usually 50 °C (47–50 °C).
Gene region | Primer name | F/R | Primer sequence 5'–3' | PCR annealing temperature (°C) | PCR/ Sequencing | Reference |
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SymF1 | F | TTTCAACWAATCATAAARAYATTGG | 49 | PCR, seq | ( |
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F | AAATGATTATTYTCWACWAATCAYAA | 50 | PCR, seq | This study |
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F | GGAGGATTTGGAAAYTGAYTAGTWCC | 49 | PCR, seq | ( |
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F | GGAGCNCCTGATATAGCWTTYCC | 47 | seq | ( |
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R | TAAACTTCWGGRTGICCAAARAATC | 47 | PCR, seq | ( |
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SymR2 | R | TAAACTTCTGGRTGTCCAAARAATCA | 47 | PCR, seq | ( |
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R | GCTCCTATTGATARWACATARTGRAAATG | 49 | PCR, seq | ( |
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F | CTYAGCCAYGCRAARGCRAARGA | 59 | PCR, seq | ( |
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F | GCWTTYTTCTCNACSAAYGCSGTNGARGG | 55 | PCR, seq | ( |
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R | TGRATRAARTGRTGRATYTCYTTIGC | 54 | PCR, seq | ( |
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NaK_910R | R | TGRATRAARTGRTGRATYTCYTT | 50 | PCR, seq | ( |
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NaK_1250Fi | F | ATGTGGTTYGAYAAYCARATYATIGA | 56 | PCR, seq | ( |
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F | ATGTGGTTYGAYAAYCARATHATIGA | 56 | PCR, seq | This study |
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R | TCGATRATYTGRTTRTCRAACCACAT | 56 | seq | ( |
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R | ACYTGRTAYTTGTTNGTNGARTTRAA | 52 | PCR, seq | ( |
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R | GATTTGGCAATNGCTTTGGCAGTDAT | 59 | PCR, seq | ( |
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POL2_104Fi | F | GYATGTCAGTYACNGATGGIGG | 59 | PCR, seq | ( |
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F | CGNATGTCNGTNACNGAYGGIGG | 60 | PCR, seq | ( |
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R | TCYTCRTTNACRTGYTTCCAYTCNGC | 59 | seq | ( |
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POL2_599F | F | GARTGGAARCAYGTVAAYGARGA | 54 | PCR, seq | ( |
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F | ATGTAYGGNTCNGCNAARAAYCARGA | 58 | PCR, seq | ( |
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POL2_889R | R | TGRAAYTGYARCATYTTWATRTTYTC | 52 | PCR, seq | ( |
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R | GGCATNCCNGGCATRTCRTTRTCNAC | 59 | PCR, seq | ( |
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F | CAYAARATGAGTATGATGGG | 51 | PCR, seq | ( |
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POL2_1459R | R | TTCATYTCRTCNCCRTCRAARTC | 52 | PCR, seq | ( |
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F | TGGGAYGGNAARATGCCNCARCC | 60 | PCR, seq | ( |
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R | GARAADATYTGYTTNCCNGTCCA | 55 | PCR, seq | This study |
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POL2_1759R | R | ATCATRTTNACRTTNCCNGGDATDAT | 55 | PCR, seq | ( |
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POL2_1777Ri | R | GTRCTGTGIGTYCKDATCATRTT | 55 | PCR, seq | ( |
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F | ACNCACAGYACNCAYCCN |
56 | seq | ( |
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POL2_2423F | F | CATTTYATHAARGAYGAYTAYGG | 51 | seq | ( |
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POL2_2509R | R | TTNACRGCRGTATCRATNAGACCYTC | 60 | PCR, seq | ( |
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R | TGNACCATNACNGAYTCCATAGCYTTDAT | 60 | PCR, seq | This study |
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POL2_2725R | R | GGATCRAAYTTRAAYTTYTTYTC | 50 | PCR, seq | ( |
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F | GYAAATTYTTYGTTGGNGGIAA | 52 | PCR, seq | ( |
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F | GTRATYGCNTGYATYGGIGARA | 52 | seq | ( |
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R | GCCCANACNGGYTCRTAIGC | 56 | seq | ( |
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R | ACNATYTGTACRAARTCWGGYTT | 52 | PCR, seq | ( |
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F | AAYAARGARGTNTTYGTNGAYTTYATGGG | 58 | PCR, seq | This study |
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F | CARTCNAARCARTTYCARCCNAARGARAC | 60 | seq | This study |
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TRRAP_1702R | R | GGNGGNCCDATNGTRTARATRTC | 56 | seq | This study |
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R | AADATYTCYTGRAANGTYTGNGGRTTCAT | 59 | seq | This study |
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F | ATGATGATHGARCCNCARAARYTNGAITA | 58 | PCR, seq | This study |
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R | TGNGCDATNGCNACCATNGTRTARTG | 60 | PCR, seq | This study |
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F | GTNTCNAAYGGNGCHATHGAYATGGCIAA | 62 | seq | This study |
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R | ACYTCYTTRTGNGGYTCCATNACYTCIGT | 62 | PCR, seq | This study |
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TRRAP_4086F | F | CARGARGCNGCNTTYGARTGYATG | 59 | seq | This study |
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R | CTRAANGTRCTNGGRAANARYTGIGT | 56 | PCR, seq | This study |
PCR reactions were carried out in a total volume of 15–35 μl containing 1.0–2.5 μl of extracted DNA, 1.5–3.5 μl (5.0–15 pmol) of primers and 7.5–17.5 μl of 2× Multiplex PCR Plus Master mix (QIAGEN). The PCR protocol consisted of an initial DNA polymerase (HotStar Taq) activation step at 95 °C for 5 min, followed by 38–40 cycles of 30 s at 95 °C, 90–120 s at 49–60 °C (depending on the primer set used), and 70–180 s (depending on the amplicon size) at 72 °C; the last cycle was followed by a final 30 min extension step at 68 °C.
Previous taxonomic publications have mostly recognised several tribes within the
Maximum likelihood tree of
Genera and species represented in Fennoscandia are marked with an asterisk (*). Species numbers are for the West Palaearctic realm, followed by Fennoscandia.
1 |
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– | * |
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2(1) |
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– |
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3(2) |
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– |
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4(3) |
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– |
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5(4,18) | * |
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– |
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6(5) | * |
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– |
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7(6) |
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– |
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8(7) | * |
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– | * |
Generic characters of
Generic characters of
Generic characters of
Preliminarily, the European
A | ( |
* |
– | ( |
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B(A) | ( |
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– | ( |
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C(B) | ( |
* |
– | ( |
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D(C) | ( |
* |
– | ( |
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E(D) | ( |
* |
– | ( |
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F(B) | ( |
* |
– | ( |
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9(4) |
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– |
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10(9) | * |
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– | [ |
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11(9) | * |
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– | * |
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12(2) | * |
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– | * |
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13(3) |
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– |
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14(13) | * |
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– | * |
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15(13) |
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– | * |
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16(7) |
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– | * |
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17(15) |
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– |
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18(17) | * |
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– |
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Generic characters of
Generic characters of
Numbers of setae on dorsal annulets are for only one side of the body, as in
1 |
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– |
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2(1) | [ |
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– |
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3(1) | ||
– |
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4(3) |
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– |
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5(4) |
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– |
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6(5) | ||
– |
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7(6) | ||
– |
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8(7) |
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– |
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9(8) | ||
– | ||
10(8) |
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– |
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11(10) |
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– |
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12(11) |
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– |
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13(12) |
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– |
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14(12) |
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– |
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15(5) | ||
– |
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16(15) | ||
– |
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17(16) | [ |
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– |
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18(17) |
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– |
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19(18) |
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20(19) |
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– |
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21(20) |
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– |
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22(17) |
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– |
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23(22) | ||
– | ||
24(4) |
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– |
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25(24) |
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– |
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26(25) |
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– |
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27(26) |
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28(26) | ||
– |
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29(25) | ||
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30(29) | ||
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31(30) | ||
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32(31) |
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33(24) |
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34(33) |
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35(33) |
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36(35) |
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37(36) | ||
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38(37) |
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39(38) |
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40(35) |
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41(40) |
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42(40) | ||
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43(42) |
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44(43) |
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45(43) |
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46(42) | ||
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Larvae of
Larvae of
Larvae of
Synonymy of genus-group names was given by
No reliable key or species treatments are available to date.
This genus was erected for a single species,
No reliable key or species treatments are available to date.
See key and species treatments in
See species treatment in
1 |
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– |
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2 | * |
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– | * |
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3 | * |
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– | * |
– lower surface of antenna noticeably paler than the upper [uniformly dark]
– medial emargination of clypeus deep, usually exceeding half of its length [reaching half of its length]
– intercostal and lanceolate cells of the fore wing and main half of the hind wing are clearly darkened [wings nearly completely hyaline]
– the 2nd anal cell of the posterior wing is almost equal to the length of the median cells [2nd anal cell of the posterior wing noticeably shorter than median one]
– 9th tergum predominantly dark [9th tergum red]
– cerci yellow [cerci basally yellow, apically fuscous]
– valvula 3 of ovipositor on lower margin noticeably convex in lateral view [only slightly convex]
– teeth of the proximal half of the ovipositor have two or more smaller additional denticles at the base [these teeth with only one small additional tooth]
Only a single paratype of
Previously published descriptions of the male of
Body length: female 6.5–8.5 mm, male 6.0–6.5 mm. Wing colour highly variable in both sexes, from nearly entirely hyaline, to entire hind wing and basal fore wing up to about pterostigma conspicuously darkened. Female (Figs
Our characterisation of the male of
See key, and notes on male (above, and under
Host plants (in Europe):
Trans-palaearctic from the British Isles, through north and central Europe (
Published records: Skåne (
Czech Republic: 1♀ (
Body length: female 5.5–8.5 mm, male 5.5 mm (only one examined). Female (Figs
We have only examined one old male specimen (DEI-GISHym31838), without genetic data, which we think belongs to
See key and notes on
Host plants:
Found widely in the Holarctic, from the British Isles, through central and northern Europe (
Published records: Skåne (
Canada: Quebec: 1♀ (DEI-GISHym15340), Gatineau Park 1.8km N Eardley, Juniperus virginiana stand, 60–80 m, 45.56667N, 76.09139W, 31.08.–07.09.2012, leg. CNC
See key and species treatments in
Only two species are known from the West Palaearctic (
1 |
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– |
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2(1) | * |
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– | * |
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3(1) | ||
– |
Body length: female 5.5–9.0 mm, male 6.5–8.0 mm. Female (Fig.
In the West Palaearctic, only
Host plants:
Widespread in central and southern Europe, from the British Isles, north to Finland (
Based on published records: Skåne, Småland (
Bulgaria: 10♀ (
No reliable key or species treatments are available to date.
No reliable key or species treatments are available to date.
Currently, fewer than 20 European taxa are considered to be
Body stocky, similar to
This is based on a translation of
Female: upper half of mesepisternum pale, lower half black. Pronotum, mesepimeron, and metapleura entirely pale. Propleuron edged with black. Head behind eyes subparallel. Antennomere 8 approx. three times as long as wide. Lancet: Fig.
Male: mesepisternum completely black. Pronotum ventrally black. Mesepimeron and metapleura partly pale. Propleuron completely black. Anterior of abdominal tergum 2 also black. Fore wing length 6.5 mm; antennomere 8 3.5 times as long as wide; head behind the eyes clearly narrowed; tergite 8 without special structures; subgenital plate apically rounded. Penis valve: Fig.
In the West Palaearctic,
Unknown.
Only known from Germany, Switzerland (
No records.
(to the best of our knowledge, the following are the only known extant collection specimens of this species):
Czech Republic [not examined: data from
One species,
France: Holotype ♀ (DEI-GISHym20818), Lorraine, Saint-Maurice-sur-Moselle, 26.05.1995, leg. Bernard (
Two species have been considered to be represented in the West Palaearctic fauna (
W. Heitland, H. Pschorn-Walcher and J. Herbst studied European populations of
Body length: female 5.0–7.0 mm, male 4.5–6.0 mm. Female: head black except for palps, and more or less labrum, underside of antennal flagellum, and sometimes more or less scape and pedicel. Thorax black, except for yellow tegula and more or less posteriodorsal edges of pronotum. Sometimes lateral edges of median mesoscutal lobe, and upper mesepisternum pale. Legs pale (orange), with dark metatarsus and apex of metatibia, and more or less dark bases of coxae. Wing venation mostly brown, with centre of fore wing
If the nearly complete loop formed by the curved up base of fore wing vein 2A+3A in
Host plants:
Widespread in Europe, from the British Isles to the Balkans, and north to Norway and Finland (
Published records:
Estonia: 3♀, 1♂ (
As already suggested by
A single larva was obtained in northern Sweden by combing through the leaves of an isolated clump of
Sweden: Torne Lappmark: 1 larva (DEI-GISHym83704), from
See
1 | * |
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— |
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Body length: female 4.5–7.5 mm, male 4.5–6.0 mm. Head black, except for mandibles and palpi. Pronotum completely black, or only extreme upper and rear edges brown. Mesepisternum more densely pubescent above than below but usually without entirely glabrous area on lower half. Hind coxa with at least basal half black. Trochanters and femora completely pale (yellowish). Tibia more whitish: pro- and mesotibia and pro- and mesobasitarsus entirely pale, with rest of tarsus darkened. Metatibia with approx. apical third black but spurs pale. Metatarsus black. Wing membrane hyaline; venation largely pale except for dark fore wing
When the shape of the claw is overlooked,
Host plants: mainly
Found through much of continental Europe, from the Iberian Peninsula and Balkans, to Finland and Norway, and also the British mainland (
Published records: Skåne (
France: 2♀ (
The conclusions on the phylogeny of
Although the
As noted above, the striking abundance and species diversity of nematine sawflies in the northern parts of the Palaearctic, including Fennoscandia, results mainly from the presence of numerous species of
As can be seen from the key to larvae, the larvae of
Although the genera which we have treated in this paper are comparatively species-poor, cases nevertheless occur of the sort of taxonomic problems which are regularly encountered in the much larger genera
This work was made possible through funding by the Swedish Taxonomy Initiative. For the loan of material, and access to collections, we thank Kees van Achterberg (