A new species of the paper wasp genus Polistes (Hymenoptera, Vespidae, Polistinae) in Europe revealed by morphometrics and molecular analyses Author Neumeyer, Rainer Author Baur, Hannes Author Guex, Gaston-Denis Author Praz, Christophe text ZooKeys 2014 400 67 118 http://dx.doi.org/10.3897/zookeys.400.6611 journal article http://dx.doi.org/10.3897/zookeys.400.6611 1313-2970-400-67 91DC4784F49A4353B4C8DC0F67B1EF92 Polistes gallicus (Linnaeus, 1767) Vespa gallica Linnaeus, 1767: 949 - Holotype male (LSL), type locality "Europa australi" [South of Europe] (not examined) Polistes foederatus Kohl, 1898: 90 - Lectotype male (NHMW), designated by Bluethgen (1943 : 129), type locality Goeygoel ["Transkauk., Helenendorf"], Azerbaijan (examined by RN) Polistes gallicus mongolicus Buysson, 1911: 218 - Syntypes males females (MNHN, ZIN), type locality road from Kuqa [ "Koutchar" ] to Karashahr [ "Karachar" ], China (Xinjiang autonomous region) (syntype male in MHNG examined by RN) Polistes foederatus obscuricornis Mader, 1936: 263 - Syntypes females (NHMW), type locality (island of) Krk, Croatia (2 syntypes examined by RN) Polistula omissa Weyrauch, 1938: 277 - Lectotype male (lost; see Arens 2011 : 462), designated by Weyrauch 1939 : 161, type locality Marseille, France, mentioned in Weyrauch (1939) : 161. Polistes omissus ordubadensis Zirngiebl, 1955: 381 - Holotype female (ZSM), type locality Ordubad, Azerbaijan (examined by RN) Polistes omissus kaszabi Giordani Soika, 1970: 327-328 - Holotype female (HNHM), type locality "Duusch ul" near Zueuenkharaa [ "Zuun-Chara" ], Mongolia (examined by RN) Polistes foederatus albellus Giordani Soika, 1976: 272 - Holotype female (HNHM, currently loaned elsewhere), type locality Bulgan aimag: Namnan ul mountains, 23 km NW of Somon Chutag, Mongolia (1 paratype female in MSNV examined by RN) Diagnosis. Relatively small and bright species with flagellum bright yellow to orange on upper side in both sexes; pedicel and extreme base of flagellomere 1 always black on upper side. Females: Epicnemial carina distinct or reduced. Hypopygium black. Metacoxa frequently spotted yellow on upper side. Mesoscutum often with pair of yellow spots. Propodeum laterally with yellow spot on each side. Clypeus breadth: malar space 3.49-4.25; head breadth: malar space 8.23-10.08; malar space: lateral ocelli distance 0.87-1.2; metatibia length: malar space 8.26-11.88; terminal flagellomere length: malar space 0.84-1.07. Males: Gena in dorsal view immediately narrowing behind eye. Epicnemium and mesosternum yellow. Head breadth: head height 1.17-1.27; lower face: clypeus breadth 1.46-1.6; terminal flagellomere length: lateral ocelli distance 1.09-1.55; terminal flagellomere length: malar space 1.28-2.21; terminal flagellomere length: terminal flagellomer breadth 1.84-3.2. Comments. The holotype of Vespa gallica Linnaeus, 1767 (LINN 2790), presently held at the Linnean Society of London, is not available for loan. We have, however, examined pictures (http://linnean-online.org/16757/). They clearly show the bright flagellum all around, excluding identity with Polistes helveticus sp. n. A careful examination of this specimen would be needed to confirm the identify of Polistes gallicus . Except for Vespa gallica and for the lost lectotype of Polistula omissa Weyrauch, 1938 we have examined and measured type specimens of all taxa listed as synonymous with Polistes gallicus . Since each of them appears to be clearly distinct from Polistes bischoffi in our morphometric analyses (Fig. 3), we only compare them with Polistes helveticus sp. n. in the following section. Bluethgen (1941: 245) and Guiglia (1972 : 49) claim that the upper side of the flagellum of Polistes foederatus Kohl, 1898 is "slightly blackened [leicht geschwaerzt]" or "darkened [assombrie]", respectively, even in the male sex, unlike the flagellum of Polistes omissus (Weyrauch, 1938). However, the flagellum of the male lectotype of Polistes foederatus from Azerbaijan is bright yellow all around, as noted by Bluethgen (1943 : 129), thus excluding any confusion with Polistes helveticus sp. n. Furthermore, the clypeus of this lectotype has not even a trace of a longitudinal furrow, although this trait has also been regarded as diagnostic for Polistes foederatus ( Bluethgen 1941: 245). Currently, both Polistes foederatus and Polistes omissus are synonyms of Polistes gallicus ( Day 1979 : 63; Gusenleitner 1985 : 105). This view is supported by our morphometric analysis (Fig. 3b, d [J]) for Polistes foederatus . The lectotype of Polistes gallicus mongolicus Buysson, 1911 has its epicnemium and mesosternum largely black as in Polistes hellenicus , but is otherwise a large, very light colored male with an extremely broad head. Its flagellum is bright all around, excluding any synonymy with Polistes helveticus sp. n. The terminal flagellomere is in fact "very short" [ tres court], as Buysson (1911 : 218) states. Morphologically it appears doubtful that the taxon Polistes mongolicus belongs to Polistes gallicus (C. van Achterberg, pers. comm.). This view is also supported by our morphometric analyses (Fig. 3b, d [K]). We have seen two syntypes (RN0444, RN0445) of Polistes foederatus obscuricornis Mader, 1936. In contrast to the statement of Mader (1936 : 263) the flagella of these two females are not "entirely black [ganz schwarz]" dorsally, but only grey. Only the scape, the pedicel, and the very base of the first flagellomere are entirely black dorsally. Both of these otherwise light colored individuals have a pronounced epicnemial carina, excluding confusion with Polistes helveticus sp. n. Although this taxon belongs without any doubt to the gallicus-group, our morphometric analyses (Fig. 3a, c [D]) do not entirely support their synonymy with Polistes gallicus . The female holotype of Polistes omissus ordubadensis Zirngiebl, 1955 is colored very light with the clypeus unspotted, the flagellum colored light dorsally, the hypopygium ( Bluethgen 1956 : 85) and even the malar space largely yellow, and the epicnemial carina distinct, all of them excluding confusion with Polistes helveticus sp. n. This view is also supported by morphometry (Fig. 3a, c [E]). In our opinion this taxon ( Polistes omissus ordubadensis ) may not even belong to the gallicus-group. The holotype of Polistes omissus kaszabi Giordani Soika, 1970 is a large, dark female with both malar space and mandibles entirely black. The epicnemial carina is distinct, excluding confusion with Polistes helveticus sp. n. Flagellum, clypeus and hypopygium are colored and patterned as in Polistes biglumis or Polistes nimpha . In fact, a synonymy with Polistes gallicus is not supported by our morphometric analyses (Fig. 3a, c [F]) and this taxon ( Polistes omissus kaszabi ) may not even belong to the gallicus-group (C. van Achterberg, pers. comm.). The holotype of Polistes foederatus albellus Giordani Soika, 1976 is currently on loan and could not be examined, but we have examined the only paratype (RN0326, MSNV-04702) mentioned in Giordani Soika (1976 : 272). It is an extremely dark female with the flagellum dorsally black and the epicnemial carina unilaterally reduced. It is morphologically and even morphometrically (Fig. 3a, c [G]) similar to a very dark Polistes helveticus sp. n., except that the bright spots and stripes are not only reduced, but also of ivory-white rather than of yellow color. These striking characters make it unlikely that this taxon ( Polistes albellus ) belongs to a species reaching Europe (C. van Achterberg, pers. comm.). These observations indicate that the taxon Polistes gallicus should be carefully revised; our analysis of the mitochondrial marker further indicates that two specimens ( Polistes sp. aff. gallicus; RN0126, RN0129) from Greece, both identified as Polistes gallicus by Arens (2011) , may represent another species. The upper side of the flagellum in both of these individuals is slightly darkened. This trait, as well as others (e.g. a broad malar space), suggest that this taxon ( Polistes sp. aff. gallicus) may be Polistes foederatus obscuricornis . More data are required to resolve this problem. In addition, our morphometric analysis reveals considerable intraspecific heterogeneity within Polistes gallicus . Specimens from continental Europe (Croatia, Portugal, Switzerland) were quite different from those from Sardinia. Both taxa overlapped but exhibited considerable differences. However, our molecular analyses did not indicate any difference between continental and Sardinian specimens. Therefore, we must also consider some unknown external factors such as temperature or humidity to be at least partially responsible for the observed morphometric variation. Such factors have previously been implicated in color pattern variation ( Zimmermann 1931 ). Material examined. 1 ♂ (Lectotype of Polistes foederatus ): Azerbaijan, Goygol District, Goeygoel ["Transkauk., Helenendorf"], 1886, Franz Friedrich Kohl det., NHMW coll.; 1 ♀ (Holotype of Polistes omissus ordubadensis : ZSM-HYM-000005): Nakhchivan Autonomous Republic, Ordubad, 1913, Kulzer leg., ZSM coll.; 1 ♂ (Lectotype of Polistes gallicus mongolicus ; EY8898): China, Xinjiang, road from Kuqa [ "Koutchar" ] to Karashahr [ "Karachar" ] in Kashgar prefecture [ "Kachgarie" ], 1909, L. Vaillant leg., MNHN coll.; 1 ♀ (RN0099): Croatia, Istria, Rovinj, Cesta za Mondelaco, 45°05'57.6"N , 13°38'36.8"E , 15 m, 24 Jul 2012, fallow, Rainer Neumeyer leg., CSE coll.; 1 ♀ (RN0100): 28 Jul 2012, fallow, Rainer Neumeyer leg., CSE coll.; 3 ♀ (RN0194, RN0195, RN0196): Rainer Neumeyer leg., RN coll.; 1 ♀ (RN0009): Rovinj, Farma Haber, 45°06'40.8"N , 13°40'21.0"E , 30 m, 22 Jul 2011, farm, Rainer Neumeyer leg., RN coll.; 1 ♀ (RN0197): Vodnjan, street D21, 44°57'58.0"N , 13°50'47.4"E , 133 m, 24 Jul 2012, fallow, Rainer Neumeyer leg., RN coll.; 2 ♀ (Syntypes of Polistes foederatus obscuricornis ; RN0444, RN0445): Krk, ≤ 1936, NHMW coll.; 1 ♀ (RN0343): Italy, Sardinia, Cabras, Stagno di Cabras, 39°57'07.6"N , 08°31'17.2"E , 5 m, 31 Aug 2013, paddy, Rainer Neumeyer leg., RN coll.; 1 ♂ (RN0347): Strada Provinciale 1, 39°55'14.1"N , 08°31'17.4"E , 6 m, 01 Sep 2013, reed, Rainer Neumeyer leg., RN coll.; 1 ♀ (RN0344): Macomer, Via Alagon, 40°15'49.9"N , 08°46'54.5"E , 553 m, 01 Sep 2013, rock face, Rainer Neumeyer leg., RN coll.; 1 ♀ (RN0345) + 1 ♂ (RN0348): Scano di Montiferro, Street SP78, 40°13'19.9"N , 08°36'29.9"E , 442 m, 01 Sep 2013, quarry, Rainer Neumeyer leg., NHM coll.; 1 ♂ (RN0346): Tadasuni, Street to San Serafino, 40°05'41.4"N , 08°52'44.5"E , 138 m, 30 Aug 2013, edge community, Rainer Neumeyer leg., RN coll.; 1 ♀ (Paratype of Polistes foederatus albellus ; RN0326, MSNV-04702): Mongolia, Bulgan aimag, Namnan ul mountains, 1150 m, 21 Jul 1968, Zoltan Kaszab leg., Antonio Giordani Soika det., MSNV coll.; 1 ♀ (Holotype of Polistes omissus kaszabi ; HNHM-283): Selenge aimag, "Duusch ul" near Zueuenkharaa [ "Zuun-Chara" ], 1100 m, 08 Jul 1964, Zoltan Kaszab leg., HNHM coll.; 1 ♀ (RN0108): Portugal, Algarve, Vila do Bispo, western shore, 37°06'57.2"N , 08°55'37.2"W , 11 m, 04 Apr 2012, dune, Rainer Neumeyer leg., RN coll.; 1 ♀ (RN0395) + 1 ♂ (RN0396): Slovakia, Nitra Region, Sturovo , 16 Sep 1947, Augustin Hoffer leg., LD coll.; 1 ♂ (RN0172): SWITZERLAND, CANTON Geneva, Geneve ,<1900, anon. leg., MHNG coll.; 1 ♂ (RN0208): CANTON Grisons, Roveredo, 18 Aug 1924, Adolf Nadig leg., ETHZ coll.; 3 ♂ (RN0205, RN0206, RN0207): 17 Aug 1949, Adolf Nadig leg., ETHZ coll.; 1 ♀ (RN0118): San Vittore, Ai Tecc, 46°14'10.42"N , 09°05'20.18"E , 262 m, 30 Jul 2012, industrial fallow, Rainer Neumeyer leg., RN coll.; 1 ♂ (RN0166): Savognin, Jul 1907, anon. leg., BNM coll.; 1 ♀ (RN0165): Val Muestair , Sta. Maria, 20 Jul 1951, Jacques de Beaumont leg., BNM coll.; 10 ♂ (RN0173, RN0209, RN0210, RN0211, RN0212, RN0213, RN0214, RN0215, RN0216, RN0217): CANTON Ticino, Locarno, 05 Jul 1907, anon. leg., ETHZ coll.; 3 ♀ (RN0110, RN0111, RN0112): Sant' Antonino, Via Gorelle/Canvera, 46°09'19.72"N , 08°58'18.65"E , 215 m, 30 Jul 2012, industrial fallow, Rainer Neumeyer leg., RN coll.; 4 ♀ (RN0103, RN0106, RN0107, RN0109): 08 Aug 2012, industrial fallow, Rainer Neumeyer leg., RN coll.; 2 ♀ (RN0041, RN0042): CANTON VALAIS, Leuk, Satellitenbodenstation, 46°19'06.66"N , 07°38'36.56"E , 919 m, 22 Aug 2011, tall herbaceous vegetation, Rainer Neumeyer leg., RN coll.; 2 ♀ (RN0104, RN0113): 07 Aug 2012, Rainer Neumeyer leg., RN coll.; 1 ♀ + 2 ♂ (RN0115, RN0116, RN0117): CANTON Vaud, Villars-sous-Yens, Arborex, 46°30'11.03"N , 06°25'12.11"E , 510 m, 25 Aug 2010, Christian Monnerat leg., MHNN coll.