(C) 2011 Marko Prous. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
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The following eleven Empria species are reported from Japan: Empria candidata (Fallén, 1808), Empria japonica Heidemaa & Prous, 2011, Empria liturata (Gmelin, 1790), Empria loktini Ermolenko, 1971, Empria plana (Jakowlew, 1891), Empria quadrimaculata Takeuchi, 1952, Empria rubicola Ermolenko, 1971, Empria tridens (Konow, 1896), Empria tridentis Lee & Ryu, 1996, Empria honshuana Prous & Heidemaa, sp. n., and Empria takeuchii Prous & Heidemaa, sp. n. The lectotypes of Poecilosoma pallipes Matsumura, 1912, Empria itelmena Malaise, 1931, Tenthredo candidata Fallén, 1808, and Tenthredo (Poecilostoma) hybrida Erichson, 1851 are designated. Empria itelmena Malaise, 1931, syn. n. is synonymized with Empria plana (Jakowlew, 1891). Poecilosoma pallipes Matsumura, 1912, previously assigned to Empria, is transferred to Monsoma, creating Monsoma pallipes (Matsumura, 1912), comb. n. Results of phylogenetic analyses using mitochondrial (COI) and nuclear (ITS1 and ITS2) sequences are also provided.
Sawflies, new species, new synonymy, key, cytochrome c oxidase I, internal transcribed spacer
With 51 valid species-level taxa (
No attempts to reconstruct the phylogeny of Empria have been made so far. Some preliminary results based on a limited number of species can be found in
Pinned specimens studied are from the following institutional collections:
BMNH Natural History Museum, London, United Kingdom (G. Broad, N. Dale-Skey Papilloud, S. Ryder, N. Springate);
CSCS Central South University of Forestry and Technology, Changsha, China (M.-C. Wei);
DEI Senckenberg Deutsches Entomologisches Institut, Müncheberg, Germany (A. Taeger, S. M. Blank, A. D. Liston);
EIHU Hokkaido University, Sapporo, Japan (M. Suwa);
HNHM Hungarian Natural History Museum, Budapest, Hungary (S. Csősz, L. Zombori);
NHRS Naturhistoriska Riksmuseet, Stockholm, Sweden (H. Vårdal);
NSMT National Museum of Nature and Science, Tokyo, Japan (A. Shinohara);
SIZ I. I. Schmalhausen Institute of Zoology, National Academy of Sciences of Ukraine, Kiev, Ukraine (I. N. Pavlusenko);
TUZ Zoological Museum of the University of Tartu, Estonia (J. Luig);
UOPJ Osaka Prefecture University, Sakai, Japan (T. Hirowatari);
USNM National Museum of Natural History, Smithsonian Institution, Washington DC, USA (D. R. Smith);
UUZM Uppsala University, Museum of Evolution, Uppsala, Sweden (H. Mejlon);
YUIC Yeungnam University Insect Collections, Gyeongsan, South-Korea (J.-W. Lee);
ZISP Zoological Institute of the Russian Academy of Sciences, St. Petersburg, Russia (S. Belokobylskij, A. Zinovjev);
ZMH Zoological Museum, Helsinki, Finland (P. Malinen);
ZML Museum of Zoology and Entomology, Lund University, Lund, Sweden (R. Danielsson);
ZMUC Zoological Museum of the University, Copenhagen, Denmark (L. Vilhelmsen).
Specimens from the private collections of Erik Heibo, Guy T. Knight, and of the second author (MH) were also studied.
For morphological analyses, male penis valves, female lancets (valvula 1), and external characters of the adults were studied.
To dissect the penis valves, genital capsules were separated from the specimen and macerated in KOH or NaOH (10–15%) for 6–12 hours at room temperature, or treated with proteinase K using High Pure PCR Template Preparation Kit (Roche, Mannheim) and following manufacturer's protocol.
Imaging methods are described in
Morphological terminology follows
For molecular phylogenetic analyses, DNA sequences of the internal transcribed spacers 1 and 2 (ITS1 and ITS2), and a mitochondrial DNA (mtDNA) fragment containing tRNA-Cys, tRNA-Tyr, cytochrome c oxidase I (COI), and partial tRNA-Leu, were obtained using methods described in
Boundaries of the sequenced tRNA and ITS2 genes were identified as described by
Phylogenetic analysis of the mitochondrial genes and combined analysis of the nuclear and mitochondrial genes were performed with MrBayes 3.1.2 (
Distances measured on the head capsule. 1 Empria quadrimaculata, head in dorsal view, female (NSMT083) (A, head length, B, head breadth, C, breadth of the postocellar area, D, length of the postocellar area, E, minimal distance between the eye and the occipital carina = head length behind the eye, F, length of the eye) 2 Empria quadrimaculata, head in anterior view, female (NSMT083) (A, minimal distance between toruli, B, malar space).
3 Monsoma pallipes, habitus in dorsal view, female (NSMT174) 4 Empria candidata, habitus in dorsal view, female (NSMT187) 5 Empria candidata, head in anterior view, female (NSMT208) 6 Empria candidata, head in dorsal view, female (NSMT208).
The data underpinning the analyses reported in this paper are deposited in the Dryad Data Repository at doi: 10.5061/dryad.fs262s48 (
1 | Abdominal terga without pale insulated (detached) paired patches (Fig. 3); length of postocellar area more than 3.5 times diameter of lateral ocellus; first flagellomere 0.9–1 times as long as flagellomeres 2–3 combined; propleura meeting broadly in front; on hind wing cross-vein m-cu present, cell M closed; valvula 1 as in Fig. 13; Hokkaido [East Palaearctic] | Monsoma pallipes |
– | Abdominal terga with pale, more or less insulated paired patches (Fig. 4); length of postocellar area less than 3.0 times diameter of lateral ocellus; first flagellomere 0.4–0.7 times as long as flagellomeres 2–3 combined; propleura not meeting or meeting only narrowly in front; on hind wing cross-vein m-cu present or absent, cell M closed or open | Empria 2 |
2 | At least facial orbits dorsally and part of temples pale (Fig. 5–6); clypeus flat without median keel; on hind wing cross-vein m-cu absent, cell M open; claws simple or with minute subbasal tooth; number of serrulae 18–21, valvula 1 as in Fig. 14; posterior margin of sternum 9 in male notched (Fig. 7), penis valve as in Fig. 25; Hokkaido [Holarctic] | Empria candidata |
– | Facial orbits and temples black (Fig. 1–2, 9–10); clypeus with median keel (distinct mostly in anterior part of clypeus only); on hind wing cross-vein m-cu usually present, cell M usually closed; claws variable; number of serrulae 13–18(19); posterior margin of sternum 9 in male rounded (Fig. 8); penis valve different | 3 |
3 | female | (female of Empria sp. 1 is currently unknown) 4 |
– | male | 13 |
4 | Postocellar area (1.9)2.1–2.5 times wider than long (Fig. 1), trochanters and trochantelli black; serrulae as in Fig. 15–16; abdominal terga with 2–3 pairs of pale patches | Empria quadrimaculata group 5 |
– | Postocellar area 1.5–2.1 times wider than long (Fig. 9–10) and / or trochanters and trochantelli pale; serrulae different (Fig. 17–24); abdominal terga with 2–6 pairs of pale patches | 6 |
5 | Abdominal terga mostly with 2 pairs of pale patches; antennae long, flagellum mostly 2.1–2.5 times longer than head breadth; in most specimens flagellomeres 1 and 2 about equally long; number of serrulae 17–19 (Fig. 15); cannot always be distinguished morphologically from Empria rubicola; Honshu, Shikoku, Kyushu | Empria quadrimaculata |
– | Abdominal terga mostly with 3 pairs of pale patches; antennae short, flagellum mostly 1.9–2.2 times longer than head breadth; in most specimens flagellomere 1 longer than flgm. 2; number of serrulae 16–18 (Fig. 16); cannot always be distinguished morphologically from Empria quadrimaculata; Hokkaido [also Sakhalin Oblast, Russia] | Empria rubicola |
6 | Malar space 2.2–2.5 times longer than lateral ocellus diameter and abdominal terga with 5–6 pairs of large pale patches; claws bifid; clypeus in most specimens at least distally pale; tegulae pale; serrulae as in Fig. 17; Hokkaido, Honshu (Yamagata) [East Palaearctic] | Empria plana |
– | Malar space 1.5–2.0 times longer than lateral ocellus diameter and abdominal terga with 2–6 pairs of small or large pale patches or malar space 1.9–2.2 times longer than lateral ocellus diameter and abdominal terga with 3 pairs of small pale patches; claws with small subbasal tooth or simple; clypeus black; tegulae black or pale; serrulae different | 7 |
7 | Serrulae as in Fig. 22–24; length of head 2.3–2.9 (2.5–3.2 in Empria tridens) times greater than length of head behind eyes (Fig. 9); trochanters and trochantelli black or slightly pale | (Empria japonica, Empria loktini, Empria tridens) 11 |
– | Serrulae as in Fig. 18–21; length of head 2.9–3.3 times greater than length of head behind eyes (Fig. 1, 10) and / or trochanters and trochantelli pale | 8 |
8 | Trochanters and trochantelli pale; tegulae completely pale | 9 |
– | Trochanters and trochantelli black; tegulae mostly black | 10 |
9 | Flagellum 2.2–2.4 times longer than breadth of head; abdominal terga with 3 pairs of small pale patches (Fig. 11); serrulae as in Fig. 18; Hokkaido, Honshu [East Palaearctic] | Empria tridentis |
– | Flagellum 1.8–2.0 times longer than breadth of head; abdominal terga with 3–4 pairs of large pale patches (Fig. 12); serrulae as in Fig. 19; Hokkaido, Honshu | Empria takeuchii |
10 | Basal serrulae conspicuously protruding (Fig. 20); claws simple or with minute subbasal tooth; abdominal terga with 5–6 pairs of pale patches; Hokkaido [Palaearctic] | Empria liturata |
– | Basal serrulae not conspicuously protruding (Fig. 21); claws with conspicuous subbasal tooth; abdominal terga with 4 pairs of pale patches; Honshu | Empria honshuana |
11 | Flagellum 2.5–2.7 times longer than breadth of head; maximal length of temple 1.40–1.55 times greater than minimal length of temple; serrulae as in Fig. 23; Hokkaido | Empria japonica |
– | Flagellum 1.8–2.3 times longer than breadth of head; maximal length of temple less than 1.35 times greater than minimal length of temple; serrulae as in Fig. 22, 24 | 12 |
12 | Abdominal terga mostly with 5 pairs of pale patches; number of serrulae 16–18 (Fig. 22); Hokkaido [Palaearctic] | Empria tridens |
– | Abdominal terga mostly with 2–3 pairs of pale patches; number of serrulae 13–14(15) (Fig. 24); Hokkaido [also Sakhalin Oblast, Russia] … Empria loktini | |
13 | Postocellar area (2.1)2.2–2.5 times wider than long and trochanters and trochantelli black; penis valves as in Fig. 26–27 | Empria quadrimaculata group 14 |
– | Postocellar area 1.7–2.1(2.2) times wider than long or trochanters and trochantelli at least partly pale; penis valves as in Fig. 28–36 | 15 |
14 | Valviceps with small basal lobe, ventroapical part of valviceps slightly bent towards its basal part (Fig. 26); flagellum 2.9–3.3 times longer than breadth of head; in most specimens flagellomere 7 not distinctly shorter than length of eye; Honshu, Shikoku, Kyushu | Empria quadrimaculata |
– | Valviceps with large basal lobe, ventroapical part of valviceps strongly bent towards its basal part (Fig. 27); flagellum 2.6–3.0 times longer than breadth of head; in most specimens flagellomere 7 distinctly shorter than length of eye; Hokkaido [also Sakhalin Oblast, Russia] | Empria rubicola |
15 | Valviceps with long apical spine (Fig. 28); malar space 1.9–2.3 times longer than lateral ocellus diameter; Hokkaido, Honshu (Yamagata) [East Palaearctic] | Empria plana |
– | Valviceps without long apical spine (Fig. 29–36); malar space 1.3–1.8 times longer than lateral ocellus diameter | 16 |
16 | Trochanters, trochantelli, and tegulae pale; abdominal terga mostly with 3 pairs of pale patches | 17 |
– | Trochanters black; trochantelli black or with barely visible median pale band or patch; tegulae black or pale; abdominal terga with 2–5 pairs of pale patches | 18 |
17 | Valviceps with large dorsobasally pointing spine at dorsoapical part (Fig. 29); postocellar area 1.9–2.3(2.4) times wider than long; flagellum 2.6–3.7 times longer than breadth of head; Hokkaido, Honshu [East Palaearctic] | Empria tridentis |
– | Valviceps with small dorsally pointing tooth at dorsoapical part (Fig. 30); postocellar area 2.0–2.7 times wider than long; flagellum 2.2–2.7 times longer than breadth of head; Hokkaido, Honshu | Empria takeuchii |
18 | Antennae short, flagellum 2.3–3.0 times longer than breadth of head | 19 |
– | Antennae long, flagellum 3.2–3.8 times longer than breadth of head | 22 |
19 | Valviceps with large dorsoapical spine (Fig. 31–32) | 20 |
– | Valviceps with small dorsoapical tooth (Fig. 33–36) | 21 |
20 | Dorsal margin of valviceps concave (Fig. 31); claws with minute subbasal tooth; abdominal terga with (2)3–4 pairs of pale patches; Honshu | Empria honshuana |
– | Dorsal margin of valviceps convex (Fig. 32); claws simple or with minute subbasal tooth; abdominal terga with 5 pairs of pale patches; Hokkaido [Palaearctic] | Empria liturata |
21 | Apical part of valvular duct extending clearly further from dorsal rim of valvura (Fig. 33); abdominal terga mostly with 2–3 pairs of pale patches; Hokkaido [also Sakhalin Oblast, Russia] | Empria loktini |
– | Apical part of valvular duct reaching almost the dorsal rim of valvura or extending only slightly further from it (Fig. 34); abdominal terga mostly with 4–5 pairs of pale patches; Hokkaido [Palaearctic] | Empria tridens |
22 | Basal lobe of valviceps short, valviceps less than 0.65 as long as valvura (Fig. 35); maximal length of temple (1.30)1.35–1.50 times greater than its minimal length; Hokkaido | Empria japonica |
– | Basal lobe of valviceps long, valviceps more than 0.8 as long as valvura (Fig. 36); maximal length of temple less than 1.35 times greater than its minimal length; Hokkaido | Empria sp. 1 |
http://species-id.net/wiki/Monsoma_pallipes
Japan, Hokkaido, Sapporo. Lectotype (here designated) female (Fig. 37), EIHU. Labelled: “Maruyama 5/24", “7", “Poecilosoma pallipes Mats., Type".
Monsoma pallipes can most easily be differentiated from the other Monsoma species, Monsoma pulveratum (Retzius, 1783), Monsoma inferentium (Norton, 1868), and Monsoma faustum Zhelochovtsev, 1961, by the colouration of the head capsule: temples, genae, facial orbits, paraantennal field laterally, and area between toruli and lateral to median ocellus are pale brown in Monsoma pallipes, while in the other three species the head capsule is black.
Unknown, but could be associated with Alnus as for Monsoma pulveratum and Monsoma inferentium (
East Palaearctic. Specimens studied are from Japan (Hokkaido) and Russia (Primorsky Krai).
Male unknown.
7 Empria candidata, posterior tip of the abdomen in ventral view, male (TUZ282970) 8 Empria quadrimaculata, posterior tip of the abdomen in ventral view, male (NSMT228) 9 Empria loktini, head in dorsal view, female (NSMT014) 10 Empria honshuana sp. n., head in dorsal view, female paratype (NSMT-Hym2011-2-3-4) 11 Empria tridentis, habitus in dorsal view, female (NSMT051) 12 Empria takeuchii sp. n., habitus in dorsal view, female paratype (NSMT032).
http://species-id.net/wiki/Empria_candidata
The morphologically closest species is the Nearctic Empria multicolor, from which Empria candidata can be distinguished by the following characters: femora predominantly and most other parts of legs at least partly black (legs are almost entirely yellowish in Empria multicolor), tarsal claws simple or with a minute inner tooth (with a long subbasal tooth in Empria multicolor), shallowly emarginated clypeus (deeply emarginated in Empria multicolor), and postocellar area more than 1.6 times wider than long (less than 1.5 in Empria multicolor) (see also
Betula (
Holarctic. Specimens studied are from China (Heilongjiang), Estonia, Finland, Japan (Hokkaido), Russia (Kamchatka Krai, Khabarovsk Krai, Leningrad Oblast, Primorsky Krai), South-Korea, Sweden, Switzerland, United Kingdom, USA (Maine).
Lancets (valvulae 1) of Monsoma and Empria. 13 Monsoma pallipes (NSMT173) 14 Empria candidata (NSMT208) 15 Empria quadrimaculata (NSMT155) 16 Empria rubicola (USNM2051678_053).
urn:lsid:zoobank.org:act:BA25596E-802D-43E3-B351-52A0BAB1B78F
http://species-id.net/wiki/Empria_japonica
USNM2051678_019: HM177347 (hologenetype COI), HM177397 (hologenetype ITS1), HM177299 (hologenetype ITS2); USNM2051678_009: HM177346 (paragenetype COI), HM177396 (paragenetype ITS1), HM177298 (paragenetype ITS2); USNM2051678_003: HM177345 (paragenetype COI), HM177395 (paragenetype ITS1), HM177297 (paragenetype ITS2).
Belongs to Empria longicornis group (see
Unknown, but could be Rubus idaeus L. subsp. melanolasius (Dieck) Focke (see
urn:lsid:zoobank.org:act:AF95BFA0-C12F-46AB-B50B-8A8CA18B34CF
Japan, Honshu, Tochigi Prefecture, Bicchuzawa, Bato, Nakagawa.
1 female, NSMT. Labelled: “[JAPAN: Honshu] Bicchuzawa, Bato, Nakagawa, Tochigi 13. IV. 2006 S. Ibuki", “NSMT110", “Holotypus ♀ Empria honshuana spec. nov. design. : M. Prous & M. Heidemaa 2011", “Empria honshuana sp.n. Prous & Heidemaa det. 2011".
“[JAPAN:Honshu] Hikagezawa Mt. Takao-san Tokyo 21. IV. 1996 A. Shinohara", 1 female, NSMT073 (NSMT); “[JAPAN: Honshu] Bicchuzawa, Bato Tochigi Pref. 9. IV. 2005 A. Shinohara" 24 males, NSMT109, NSMT115, NSMT121–137, NSMT166–170 (NSMT), 1 male TUZ615362 (TUZ); “[JAPAN: Honshu] Bicchuzawa, Bato Tochigi Pref. 23. IV. 2005 A. Shinohara" 1 male, NSMT171 (NSMT); “[JAPAN: Honshu] Bicchuzawa, Bato Tochigi Pref. 29. IV. 2005 A. Shinohara" 1 female, TUZ615361 (TUZ); “[JAPAN:Honshu] Annaigawa, nr Mt. Takao-san Tokyo 17. IV. 1994 A.&T.Shinohara" 1 female, NSMT198, 2 males, NSMT120, NSMT200 (NSMT); “[JAPAN:Honshu] Akigase-koen Saitama Pref. 14. IV. 1996 A. Ta., N. & To. Shinohara" 1 female, NSMT204 (NSMT); “[JAPAN: Honshu] Bicchuzawa, Bato Nakagawa, Tochigi 13. IV. 2006 S. Ibuki" 1 male, NSMT106 (NSMT); “[JAPAN:Honshu] Bicchuzawa Bato, Tochigi 1. V. 2010 S. Ibuki" 1 female, NSMT-Hym2011-2-3-4 (NSMT); “JAPAN: Chiba Pref. Okusa-cho, Wakaba-shi 35°36.5'N, 140°11.6E' 23 March 1997 O. S. Flint, Jr." 1 female, USNM2051678_016 (USNM); “JAPAN: Honshu Himuro-machi Utsunomyia-shi Tochiji-ken [Utsunomiya-shi Tochigi-ken], Mal. 2-15.IV.2009, Mal. trap Takeyuki Nakamura leg." 1 male, USNM2057434_04 (USNM).
NSMT106: JN029870 (paragenetype COI), JN029890 (paragenetype ITS1), JN029854 (paragenetype ITS2); NSMT-Hym2011-2-3-4: JN029891 (paragenetype ITS1); USNM2051678_016: JN029871 (paragenetype COI), JN029892 (paragenetype ITS1).
Body length. 6.0–6.9 mm.
Colour. Black; following parts unpigmented, pale: apical maxillary palpomeres; posterodorsal margin of pronotum in lateral parts; tegulae (except lateroproximal part); median band or patch of pro-, meso-, and metatrochantellus; profemur apically; protibia in anterior and partly posterior aspects; mesotibia partly in anterior and posterior aspects; metatibia basally; tarsomere 1 of hind leg basally; paired patches on abdominal terga 2–5; at least partially posterior margins of terga (tergum 10 dorsally more widely) and sterna; and cenchri. Labrum from yellowish-brown to blackish.
Head. Head behind eyes in dorsal view subparallel sided; postocellar area trapeziform, its length equal to or longer than 2 times diameter of lateral ocellus; distinct and diverging lateral postocellar furrows going from ocelli towards occiput at least to the distance of ocellus diameter; area between frontal crests clearly exceeding the level of crests in dorsal view; postocellar area with indistinct punctures and interspaces, more or less glossy; punctures more regular on temples and postocular area, face with more irregular punctures; wrinkled interspaces more prominent on frontal area; clypeus with rough irregular punctures, more or less fused; ocellar and postocellar area convex, slightly raised; clypeus tridentate with median keel distinct mostly in anterior part of clypeus only, median tooth smaller than lateral teeth; malar space about equal to or shorter than distance between antennal sockets; frontal ridge V-shaped; pit in central part of frontal field present; median ocellus surrounded by groove, with short distinct longitudinal furrow anteriorly, and with similar but mostly less distinct furrow posteriorly. Maximal length of temple 1.2–1.4 times greater than its minimal length; flagellum 1.9–2.0 times longer than breadth of head.
Thorax. Mesoscutellum, mesoscutellar appendage, and metapostnotum more or less glossy, almost impuctate or with indistinct shallow punctures; metascutellum with irregular fine punctures; punctures on mesoscutum more evident on lateral and anterior regions of the median lobes, fading towards central regions; mesepisternal punctures variable between specimens, from rather weak with intespaces almost glossy to more distinct with sculptured, interspaces; mesepimeron with setae on posterior part; metepisternum with evenly distributed setae; metepimeron in central part without setae; distance between cenchri 1.1–1.4 times of cenchrus width; wings hyaline, venation brownish, becoming paler near junction to thorax; closed cell M in hindwing present; tarsal claws with conspicuous subbasal tooth.
Abdomen. Terga on most parts with transverse keel-like sculpticells and with short setae (about half of lateral ocellus diameter), sometimes with shallow punctures at median parts of terga 2–4; posterior parts of terga (6) 7–9 (occasionally terga 3–10) at median line with small more or less triangular pale regions; ventral margin of valvula 3 slightly bending towards apex, slightly longer than valvifer 2; serrulae of valvula 1 as in Fig. 21, number of serrulae 15–16.
(Mostly the differences compared to female are given).
Body length. 4.8–5.6 mm.
Colour. Unpigmented, whitish or yellowish brown: anterolateral (seldom also posterolateral) margins of tegulae; protibia in anterior aspect, often partly also in posterior aspect; mesotibia partly in anterior aspect; outer margins of harpes; and paired patches on abdominal terga 2–(3)/4/(5).
Head. Area between frontal crests reaching or slightly exceeding the level of crests in dorsal view; malar space less than or equal to distance between antennal sockets; length of postocellar area about 2 times of lateral ocellus diameter; maximal length of temple 1.25–1.45 times greater than its minimal length; flagellum 2.3–2.6 times longer than breadth of head.
Thorax. Distance between cenchri variable, up to 2 times width of cenchrus. Tarsal claws with minute subbasal tooth.
Abdomen. Tergum 8 with indistinct tergal hollows which form semioval or semicircular depression reaching 1/3–1/2 of tergum length and sometimes possessing indefinite central procidentia. Posterior margin of sternum 9 round; penis valve as in Fig. 31.
Based on the similarities in penis valves, the closest species is Empria sulcata Wei & Nie, 1998 from China (see http://www.morphbank.net/?id=643394). While the penis valves of both species can easily be distinguished, the distinctly concave dorsal margin of valviceps of these species is a unique characteristic within Empria. Serrulae of the two species are clearly different (cf. Fig. 21 and http://www.morphbank.net/?id=700325). Externally the species can mainly be distinguished by colouration: in Empria sulcata tegulae are completely pale and legs extensively yellowish, while in Empria honshuana tegulae are at least partly and legs predominantly black.
Unknown.
Japan (Honshu).
The species name refers to the type locality, Honshu, the main island of Japan.
Lancets (valvulae 1) of Empria. 21 Empria honshuana sp. n., paratype (USNM2051678_016) 22 Empria tridens (USNM2051678_018) 23 Empria japonica, holotype (NSMT USNM2051678_019) 24 Empria loktini (TUZ615180).
http://species-id.net/wiki/Empria_liturata
The most similar species morphologically appears to be Nearctic Empria ignota (Norton, 1867). The clearest differences between these species can be seen in the structure of penis valves (Fig. 32; http://www.morphbank.net/?id=694564).
Filipendula ulmaria (L.) Maxim., Geum rivale L. (based on ex ovo rearings by MP in Estonia). Fragaria vesca has also been suggested (
Palaearctic. Specimens studied are from Belgium, Croatia, Czech Republic, Denmark, Estonia, France, Germany, Hungary, Italy, Japan (Hokkaido), Russia (Leningrad Oblast), Switzerland, United Kingdom.
Penis valves of Empria. 25 Empria candidata (NSMT036) 26 Empria quadrimaculata (UOPJ03) 27 Empria rubicola (USNM2051678_042) 28 Empria plana (NSMT201) 29 Empria tridentis (TUZ615182) 30 Empria takeuchii sp. n., paratype (NSMT112).
http://species-id.net/wiki/Empria_loktini
Belongs to Empria longicornis group, morphologically the closest is Empria basalis Lindqvist, 1968, which can be distinguished from Empria loktini by clearly different penis valves, lancets (see
Unknown.
East Palaearctic. Specimens studied are from Japan (Hokkaido) and Russia (Sakhalin Oblast).
http://species-id.net/wiki/Empria_plana
Morphologically the closest species is Empria immersa (Klug, 1818), from which Empria plana can be distinguished by differences in the structure of serrulae (Fig. 17; http://www.morphbank.net/?id=694567) and penis valves (Fig. 28; http://www.morphbank.net/?id=578888). Externally, the Empria plana specimens from mainland Asia differ clearly from Empria immersa also by their pale clypeus (black in Empria immersa), which is, however, only partly pale or nearly black in Japanese specimens. In this regard, some disagreements concerning the taxonomic status of Empria plana should also be noted. Some authors treat this taxon either as a geographical form, or as a subspecies of Empria immersa (
Possibly Salix sp., see Verzhutskii (1966; 1981)under the name Empria immersa.
East Palaearctic. Specimens studied are from Japan (Hokkaido, Honshu), Mongolia, and Russia (Amur Oblast, Irkutsk Oblast, Kamchatka Krai, Khabarovsk Krai, Primorsky Krai).
Penis valves of Empria. 31 Empria honshuana sp. n., paratype (NSMT200) 32 Empria liturata (USNM2051678_051) 33 Empria loktini (NSMT105) 34 Empria tridens (USNM2051678_024) 35 Empria japonica, paratype (NSMT009) 36 Empria sp. 1 (USNM2051678_040). The arrowheads illustrate the different position of valvular duct (upper right arrowhead) relative to the dorsal rim of valvura (lower left arrowhead) in Empria loktini (Fig. 33) and other species of longicornis-group (Fig. 34).
http://species-id.net/wiki/Empria_quadrimaculata
The closest species are Empria zhangi Wei & Yan, 2009 (China) and Empria rubicola Ermolenko, 1971. Empria zhangi (two females and two males studied, including the holotype) can be distinguished from Empria quadrimaculata mainly by the following two characters: 1) in female malar space clearly less than two times of the lateral ocellus diameter (about two times in Empria quadrimaculata and Empria rubicola), in male equal or slightly less than the ocellus diameter (clearly longer in Empria quadrimaculata and Empria rubicola); and 2) in female flagellum about 2.0 times longer than breadth of head (2.1–2.5 times in Empria quadrimaculata), in male 2.4–2.5 times (2.9–3.3 times in Empria quadrimaculata). Empria rubicola has shorter antennae and three pairs of pale patches (mostly two in Empria quadrimaculata) on terga. The penis valves of Empria zhangi and Empria quadrimaculata are very similar (http://www.morphbank.net/?id=693502; Fig. 26), while Empria rubicola can be distinguished from the two by relatively large basal lobe of the valviceps and by the ventroapical part clearly bent towards its basal part (Fig. 27). Valvula 1 appears indistinguishable in all three species.
Japan (Honshu, Shikoku, Kyushu).
http://species-id.net/wiki/Empria_rubicola
The closest species are Empria zhangi and Empria quadrimaculata (see under Empria quadrimaculata Takeuchi, 1952 for details).
Unknown. Holotype female and the studied paratypes (1 female, 2 males) were collected from Rubus idaeus L. subsp. melanolasius (Dieck) Focke(under the name Rubus sachalinensis in
East Palaearctic. Specimens studied are from Japan (Hokkaido) and Russia (Sakhalin Oblast). Most probably this species has to be removed from the list of Chinese species (
urn:lsid:zoobank.org:act:BDE02124-C81A-4705-91F4-34B40134B0C1
Japan, Honshu, Yamanashi Prefecture, Utsukushinomori, Yatsugatake Mts.
1 female, NSMT. Labelled: “[JAPAN:Honshu] Utsukushinomori 1500–1700m Yatsugatake Mts. Yamanashi Pref. 5–8. VI. 2000 A. Shinohara", “NSMT044", “Holotypus ♀ Empria takeuchii sp. n. design. : M. Prous & M. Heidemaa 2011", “Empria takeuchii sp.n. Prous & Heidemaa det. 2011".
“Shimashima Nagano Pref 16. V. 1984 A. Shinohara", 1 female, NSMT032 (NSMT); “[JAPAN:Honshu] Kamiange, Mt. Jinba Tokyo 27. IV. 2003 A. Shinohara", 1 male, NSMT037 (NSMT); “Ōmi, Ō hara [Ōhara] Kyoto Pref. 15. V. 1984 R. Inagawa", 1 female, NSMT041 (NSMT); “[Ōmi, Ōhara] Sakyo-ku, Kyoto Kyoto Pref. May, 14, 1984 T. Matsumoto leg." 1 female, NSMT211 (NSMT); “[JAPAN: Honshu] Yokotemichi, ca. 850m 35-22-39N 133-31-21E Mt. Daisen Tottori Pref. 28-29. IV. 2007 A. Shinohara“, 1 male, NSMT112 (NSMT); “Takihata Kawachi-Nagano Osaka 22. IV. 1981 A. Shinohara", 1 male, NSMT213 (NSMT); “JAPAN: Ishikawa Pref., Mt. Shiritaka 637 m, May 19 1979 D. Smith & I. Togashi" 1 female, USNM2051678_047 (USNM); “JAPAN: Honshu Tamozawa, Nikkô-shi Tochigi-ken, Mal. trap 13-27.iv.2009 Takeyuki Nakamura leg.", 1 male, USNM2057434_03 (USNM).
“JAPAN, Hokkaido Ginsendai, Kamikawa-chô 43°40'N, 143°01'E, 947 m Selectively cut forest 6–27.vi.2008 Mal. trap, A. Ueda leg" 1 female, USNM2051678_011 (USNM); “JAPAN, Hokkaido Sekihoku-tôge, Kamikawa-chô, natural forest, 993 m 43°40'N, 143°06'E, 6–27.vi.2008 Mal. trap, A. Ueda leg." 3 males, USNM2051678_008, USNM2051678_031, USNM2051678_061 (USNM); “42°57'N, 141°14'E Hakken-zan Sapporo, Hokkaidō JAPAN 16.v.2009 Takuma YOSHIDA leg." 2 males, USNM2057434_06, USNM2057434_07 (USNM).
Body length. (5.1)6.4–6.9 mm.
Colour. Black; following parts more or less unpigmented, whitish or yellowish brown: labrum; apical maxillary and labial palpomeres; tegulae completely; posterodorsal margin of pronotum in lateral part rather widely, upper part of posterolateral margin of pronotum quite narrowly; pro-, meso-, and metacoxa apically; pro-, meso-, and metatrochanter partly or in most part; pro-, meso-, and metatrochantellus partly or completely; profemur in anterior, posterior, and lateral aspects; mesofemur and metafemur apically slightly; protibia in anterior and posterior aspects; mesotibia in most part; metatibia in basal 2/3; tarsomere 1 of hind leg in basal 2/3; paired patches on abdominal terga 2–4(5); posterior margins of terga and sterna; and cenchri (in one female only posterior margin).
Head. Head behind eyes in dorsal view subparallel sided; postocellar area trapeziform, its length mostly less than or equal to 2 times of lateral ocellus diameter; area between frontal crests in dorsal view reaches or slightly exceeds the level of crests; face and clypeus with somewhat irregular punctures, less shining compared to vertex and especially to postocellar area; ocellar and postocellar area at least slightly raised; clypeus tridentate, with median tooth smaller than lateral teeth; clypeus with median keel; malar space (minimal ventro-ocular distance) shorter or equal to distance between antennal sockets; frontal ridge “V"-shaped, central part of frontal field with distinct pit; maximal length of temple 1.25–1.4 times greater than its minimal length; flagellum 1.8–2.0 times longer than breadth of head.
Thorax. Anterior part of mesoscutum with more or less distinct punctures, its median and postero-lateral portions in most part with sparse indistinct punctures and glossy interspaces, or almost impunctate, glossy; mesoscutellum, mesoscutellar appendage, and metapostnotum impunctate and glossy; mesepisternum with more or less indistinct punctures, mostly glossy; mesepimeron with setae on posterior part; metepisternum with evenly distributed setae; metepimeron in central part without setae; distance between cenchri in most specimens about equal to cenchrus width, but sometimes slightly greater; wings hyaline with brownish venation; closed cell M in hindwing present; tarsal claws with conspicuous subbasal tooth.
Abdomen. Terga mostly with keel-like (sometimes mixed with scale-like) sculpticells and short setae (about half of lateral ocellus diameter); ventral margin of valvula 3 abruptly bending towards apex, about equal in length to valvifer 2; serrulae of valvula 1 as in Fig. 19, number of serrulae (15)16–17.
(Mostly the differences compared to female are given).
Body length. 5.6–5.8 mm.
Colour. Unpigmented, whitish or yellowish are: meso- and metatrochanter apically; pro-, meso-, and metatrochantellus partly; mesofemur only apically, or in anterior, posterior, and lateral aspects; metafemur apically; mesotibia partly in anterior, posterior, and lateral aspects, or in most part; metatibia in basal 1/3 or in basal 1/2; outer margins of harpes; paired patches on abdominal terga 2–4(3).
Head. Area between frontal crests in dorsal view not exceeding the level of crests; length of postocellar area 1.5–2.0 times of lateral ocellus diameter; maximal length of temple 1.25–1.45 times greater than its minimal length; flagellum 2.2–2.7 times longer than breadth of head.
Abdomen. Posterior margin of sternum 9 round; tergum 8 without tergal hollows and procidentia; penis valve as in Fig. 30.
Morphologically, no certain closest relative can be specified. Superficially may resemble Empria rubicola (based on males), Empria honshuana (based on females), or Empria tridentis (both have pale trochanters and trochantelli). Penis valve (Fig. 30) and valvula 1 (Fig. 19) clearly distinguish this species from all other known species of Empria. According to the molecular analyses (of ITS1 and ITS2 combined with mtDNA sequences), the closest species are those of the Empria longicornis and Empria immersa species groups, and Empria tridentis (Fig. 38, 40).
Unknown.
Japan (Hokkaido, Honshu).
The specific name refers to Kichizo Takeuchi (1892–1968), who made great contributions to the sawfly systematics in eastern Asia.
Six additional studied specimens (1 female, 5 males) from Hokkaido were not included in the type series. The female and most of the males have a longer postocellar area (more than 2 times of the lateral ocellus diameter) compared to the specimens from Honshu (mostly less than 2 times). Serrulae of the Hokkaido female are also slightly different (cf. http://www.morphbank.net/?id=693521 and Fig. 19). No clear differences were found in the structure of penis valves between the specimens from Hokkaido and Honshu.
http://species-id.net/wiki/Empria_tridens
Belongs to Empria longicornis group. Morphologically the closest species is Empria longicornis, from which it can be distinguished in most cases by shorter antennae and more pairs of pale patches on abdominal terga (4 large and 1 small in Empria tridens, on terga 2–6; 3 large and 1 small in Empria longicornis, on terga 2–5), and by its less prominent serrulae (Fig. 22; http://www.morphbank.net/?id=578850).
Rubus idaeus and possibly Rubus fruticosus complex (
Palaearctic. Specimens studied are from Belgium, Croatia, Denmark, Estonia, Finland, France, Germany, Hungary, Japan (Hokkaido), Mongolia, Russia (Amur Oblast, Kamchatka Krai, Kostroma Oblast, Leningrad Oblast, Primorsky Krai, Sakhalin Oblast, Stavropol Krai, Volgograd Oblast), Sweden, Switzerland, Turkey, Ukraine, and United Kingdom.
http://species-id.net/wiki/Empria_tridentis
Morphologically, no close relatives can be identified, but in the phylogenetic analysis of the ITS and mtDNA sequences combined, the species appears as a sister of the longicornis-group (Fig. 40). Superficially may resemble Empria longicornis, from which Empria tridentis can easily be distinguished by tegulae, base of metatibia, trochanters, and trochantelli pale (all black in Empria longicornis), and by very different structure of lancets and penis valves.
Unknown.
East Palaearctic. Specimens studied are from Japan (Hokkaido, Honshu), Russia (Khabarovsk Krai, Primorsky Krai), and South-Korea.
The original description of this species states that there are “a pair of large flecks on lateral portion of lst–4th tergite" (
Belongs to Empria longicornis group. Externally it is most similar to Empria japonica, but penis valve is clearly distinct from all other known species of the longicornis-group (Fig. 36), being most similar to Empria alpina Benson, 1938 (e.g. http://www.morphbank.net/?id=577439). Can be distinguished from Empria alpina by its colouration: in Empriasp1 tegulae, posterior margin of pronotum, and basal 1/3 of metatibia are pale, while in Empria alpina these are mostly black. Distinctness of this taxon is also supported by nuclear ITS sequence data (Fig. 38).
Unknown.
Japan (Hokkaido).
Because taxonomy of the longicornis-group is quite difficult (
Bayesian analyses of the mitochondrial and nuclear sequences separately and in combination all resulted in somewhat different topologies (Fig. 38–40), with well supported differences in some cases (especially in the longicornis and the immersa-groups). However, several clades were reconstructed in all analyses with significant statistical support (posterior probability 0.95 or more). Based on these analyses, the basal split within the genus Empria is between Empria candidata and all other species (Fig. 38–40), which is consistent with the division of the genus into two subgenera, Parataxonus MacGillivray, 1908 (Empria candidata) and Empria s. str. (
Each of Empria japonica, Empria loktini, Empria longicornis, Empria immersa, and Empria plana is monophyletic (as would be expected from morphology) according to the ITS sequences (Fig. 38), but not according to the mitochondrial DNA (Fig. 39). The monophyly of Empria tridens is supported neither by ITS nor the mitochondrial sequences (Fig. 38–39; see discussion in
Monsoma pallipes, lectotype of Poecilosoma pallipes Matsumura, 1912, habitus in dorsolateral view, female.
Phylogenetic analyses of the genus Empria. 38 Phylogeny of ITS sequences (1298–1517 bp) reconstructed using BAli-Phy (GTR + I + G[4] substitution model). Because the four independent runs of BAli-Phy produced different topologies, only clades which were found in all trees and were supported with posterior probabilities (PP) 0.9 or more are shown. Duplicate (shown behind the sequence used in the analysis) and very similar sequences (three Empria japonica, two Empria tridentis, and one Empria rubicola) were removed prior to analyses to reduce computation time. 39 Phylogeny of mitochondrial sequences using MrBayes (GTR + I + G[4] model; alignment length 1642 bp). Duplicate sequences (shown behind the sequence used in the analysis) were removed prior to analyses. Empria liturata from Japan (USNM2051678_021) was also excluded due to incomplete sequence. 40 Combined analysis of ITS (MAP alignment from BAli-Phy analysis) and mitochondrial sequences using MrBayes (GTR + I + G[4] model). Monsoma pulveratum was used as an outgroup. Clades with posterior probabilities (PP) less than 0.9 were collapsed in all the trees.
Although identification of Empria species using only external morphology can often be difficult, we found that females of the species reviewed here can mostly be identified without dissecting their ovipositors. Identification of the males is much less reliable without studying their genitalia because of more extensive intraspecific variation and less pronounced differences among species. The most difficult species to separate from each other on the basis of female characters are Empria quadrimaculata and Empria rubicola, the ovipositors of which appear nearly indistinguishable (Fig. 15–16). Also the external characters applied in the present key overlap considerably between them. However, because there are consistent differences in the penis valves between the two (see Fig. 26–27), they most likely represent different species.
Due to the general difficulty in identifying the Empria species using only external morphology, it is advisable in our opinion to leave the specimens unidentified (to avoid possible confusions in the future), especially those from the poorly studied regions (e.g. Eastern and Central Asia), as long as their identity remains problematic from external morphology and the genitalia cannot be dissected.
In addition to the 11 named Empria species and one presumably new but undescribed species (currently only one male is known) reported here, some additional species of the genus are likely to be found in Japan. Alpine habitats above the tree line might be inhabited by additional Empria species, but from there we have no samples yet.
The results of our molecular phylogenetic analyses (Fig. 38–40) significantly supported the groupings within Empria that could be expected from morphology (Empria s. str., immersa-group, longicornis-group, and quadrimaculata-group). Although Empria pumiloides was the only species from the hungarica-group in the current dataset, monophyly of this group is also supported by DNA data (unpublished results). The consistent affinity found between the longicornis-group, the immersa-group, and Empria tridentis in all our analyses (Fig. 38–40) was the only phylogenetic result not expected from morphology (though phylogenetic analyses using morphological data are still lacking). Based on the phylogenetic results presented here, we cannot draw any more definite conclusions regarding the phylogeny of Empria, which require, in addition to improving taxon and gene sampling, possibly also methodological advancements (e.g. using methods which take into account incomplete lineage sorting;
We would like to thank Sergey A. Belokobylskij and Alexey G. Zinovjev (ZISP), Olof Biström and Pekka Malinen (ZMH), Stephan M. Blank (DEI), Sándor Csősz and Lajos Zombori (HNHM), Roy Danielsson (ZML), Toshiya Hirowatari (UOPJ), Jong-Wook Lee (YUIC), Andrew Liston (DEI), Ole Martin and Lars Vilhelmsen (ZMUC), Hans Mejlon (UUZM), Inna N. Pavlusenko and Valery A. Korneyev (SIZ), Suzanne Ryder, Natalie Dale-Skey Papilloud, Gavin Broad (BMNM), David R. Smith (USNM), Masaaki Suwa (EIHU), Andreas Taeger (DEI), Hege Vårdal (NHRS), and Meicai Wei (CSCS) for loaning us material (including type specimens) from institutional collections. Stephan M. Blank clarified for us localities of some of the types. Madli Pärn and Andro Truuverk are thanked for technical assistance. Kauri Mikkola (ZMH) kindly commented on the new species names. Comments and suggestions by Stephan M. Blank, Toomas Tammaru (University of Tartu) and two anonymous reviewers helped to improve the manuscript.
The study was financially supported by the Estonian Science Foundation grant nr. 6598 to MH, the Estonian Ministry of Education and Science (target-financing project number 0180122s08) and the European Union through the European Regional Development Fund (Center of Excellence FIBIR).