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. 2014 Apr 11:(400):67-118.
doi: 10.3897/zookeys.400.6611. eCollection 2014.

A new species of the paper wasp genus Polistes (Hymenoptera, Vespidae, Polistinae) in Europe revealed by morphometrics and molecular analyses

Rainer Neumeyer  1 , Hannes Baur  2 , Gaston-Denis Guex  3 , Christophe Praz  4
Affiliations

A new species of the paper wasp genus Polistes (Hymenoptera, Vespidae, Polistinae) in Europe revealed by morphometrics and molecular analyses

Rainer Neumeyer et al. Zookeys. .

Abstract

We combine multivariate ratio analysis (MRA) of body measurements and analyses of mitochondrial and nuclear data to examine the status of several species of European paper wasps (Polistes Latreille, 1802) closely related to P. gallicus. Our analyses unambiguously reveal the presence of a cryptic species in Europe, as two distinct species can be recognized in what has hitherto been considered Polistes bischoffi Weyrauch, 1937. One species is almost as light coloured as P. gallicus, and is mainly recorded from Southern Europe and Western Asia. The other species is darker and has a more northern distribution in Central Europe. Both species occur syntopically in Switzerland. Given that the lost lectotype of P. bischoffi originated from Sardinia, we selected a female of the southern species as a neotype. The northern species is described as P. helveticus sp. n. here. We also provide a redescription of P. bischoffi rev. stat. and an identification key including three more closely related species, P. biglumis, P. gallicus and P. hellenicus.

Keywords: Cytochrome oxidase 1; DNA barcoding; ITS; LDA ratio extractor; Switzerland; allometry; cryptic species; linear discriminant analysis; principal component analysis; ratio spectrum; shape PCA.

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Figures

Figure 1.
Figure 1.
Phylogenetic tree based on maximum likelihood analysis of sequences of the mitochondrial gene cytochrome oxidase 1 (COX1); numbers shown at nodes are bootstrap values based on 1000 bootstrap replicates performed in RAxML v.7.0.4. Branches interrupted by an oblique line have been shortened for better graphic representation.
Figure 2.
Figure 2.
Phylogenetic tree based on maximum likelihood analysis of sequences of the nuclear marker ITS1; numbers shown at nodes are bootstrap values based on 1000 bootstrap replicates performed in RAxML v.7.0.4. Branches interrupted by an oblique line have been shortened for better graphic representation.
Figure 3.
Figure 3.
Shape PCA of all five OTUs of the Polistes gallicus-group. a, b Scatterplot of first against second shape PC a females b males c, d Scatterplot of isosize against first shape PC c females d males. Symbols: orange circles = Polistes biglumis, purple triangles = Polistes bischoffi, green diamonds = Polistes gallicus, blue rectangles = Polistes hellenicus, red dots = Polistes helveticus. In parentheses, the variance explained by each shape PC. Encircled upper case letters show the position of the type specimens of various nominal taxa: (A) Polistes bimaculatus pamirensis Zirngiebl, 1955, syntypes; (B) Polistes bimaculatus nigrinotum Zirngiebl, 1955, holotype; (C) Polistes bischoffi Weyrauch, 1937, neotype; (D) Polistes foederatus obscuricornis Mader, 1936, syntypes; (E) Polistes omissus ordubadensis Zirngiebl, 1955, holotype; (F) Polistes omissus kaszabi Giordani Soika, 1970, holotype; (G) Polistes foederatus albellus Giordani Soika, 1976, paratype; (H) Polistes helveticus Neumeyer sp. n., holotype; (I) Polistes dubius Kohl, 1898, lectotype; (J) Polistes foederatus Kohl, 1898, holotype; (K) Polistes gallicus mongolicus Buysson, 1911, syntype; (L) Polistes hellenicus Arens, 2011, holotype.
Figure 4.
Figure 4.
Scatterplot of first shape PC against isosize for comparison of Polistes bischoffi with Polistes helveticus: a females b males. Symbols: purple triangles = Polistes bischoffi, red dots = Polistes helveticus; closed symbols = specimens identified by genetic clustering and morphological characters; open symbols = specimens identified by morphological characters only. In parentheses the variance explained by the first shape PC.
Figure 5.
Figure 5.
PCA ratio spectrum (blue) and allometry ratio spectrum (green) of a comparison of Polistes bischoffi with Polistes helveticus. a females b males. Horizontal bars in PCA and allometry ratio spectrum represent 68% bootstrap confidence intervals based on 1000 replicates.
Figure 6.
Figure 6.
Different aspects of a female (specimen RN0137) of Polistes bischoffi Weyrauch, 1937: a frontal view of head b lateral view of lower face with malar space and mandible c lateral view of head and mesosoma d lateral view of body e body from above. Arrows indicate the yellow patch on the lateral part of propodeum (c) and one of two yellow spots on the mesoscutum (e).
Figure 7.
Figure 7.
Different aspects of a male (specimen RN0151) of Polistes bischoffi Weyrauch, 1937: a frontal view of head b lateral view of lower face with malar space and mandible c dorsal view of body d lateral view of head, mesosoma, and base of metasoma e lateral view of body. The red arrow in b is pointing to the faint lateral ridge on clypeus. In d the white arrow is pointing to the yellow ventrolateral stripe of the pronotum, the red arrow to the lateral extension of terminal band on tergum 2, the black arrow to a large yellow spot on sternum 2 and the blue arrow on the yellow basal band of sternum 3.
Figure 8.
Figure 8.
Lateral aspect of the holotype (individual RN0242; male) of Polistes hellenicus Arens, 2011. Unique for males of European Polistes is the black ventral side of the mesosoma, especially epicnemium and mesosternum.
Figure 9.
Figure 9.
Different aspects of a female (individual RN0138) of Polistes helveticus sp. n.: a frontal view of head b lateral view of lower face with malar space and mandible c mesopleural region of mesosoma d lateral view of body e dorsal view of body. The red arrow in picture (c) is pointing to the rather reduced epicnemial carina, and the white arrow to the quite distinct mesopleural signum (sensu Carpenter 1996a), a structure also called a sternopleural groove (Richards 1973).
Figure 10.
Figure 10.
Different aspects of a male (individual RN0153) of Polistes helveticus sp. n.: a frontal view of head b anterolateral view of lower face c lateral view of body d dorsal view of body. The arrows are pointing to the isolated black area (a) bordering the torulus, the very faint lateral ridges of clypeus (b) or the ventrolateral angle of the pronotum (c).
Figure 11.
Figure 11.
Distribution of examined specimens of Polistes bischoffi Weyrauch, 1937 and Polistes helveticus sp. n. While Polistes bischoffi mainly occurs from Southern Europe to Western Asia, Polistes helveticus appears to have a more northern distribution in Central Europe. Thus far, the only incidences of syntopy (Polistes bischoffi, Polistes helveticus) are from Switzerland.
Figure 12.
Figure 12.
Drawings of morphological characters used in the key to European paper wasps (Polistes) of the gallicus-group: Mesopleuron with abrupt (a) or gradual (b) sculpture change; female antennae with dark (c) or bright (d) upper side of flagellomeres; male apical flagellum dark (e) or bright (f) on upper side; head in frontal view with black band across clypeus (g) or isolated black spot (h); apical mesosoma in dorsal view with drop-shaped spots on mesoscutum, posterior pronotal stripes ending far from pronotal carina (i) or without drop-shaped spots, pronotal stripes approaching pronotal carina (k); head in dorsal view with genae convex (l) or immediately narrowing (m). Drawings a, b, c, d, f are courtesy of H. Cigler, g, h, i, k of D. Lawniczak, and e, l, m of CSCF.
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