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(3) Baracchi.Indd 191 03-08-11 09:29 192 Baracchi Et Al Contributions to Zoology, 80 (3) 191-199 (2011) Relevance of wing morphology in distinguishing and classifying genera and species of Stenogastrinae wasps David Baracchi1, 2, Leonardo Dapporto1, Stefano Turillazzi1 1 Università degli Studi di Firenze, Dipartimento di Biologia Evoluzionistica ‘Leo Pardi’, Via Romana 17, 50125, Firenze, Italy 2 E-mail: [email protected] Key words: geometric morphometrics, Hymenoptera, nest architecture, taxonomy, Vespidae Abstract to the generalized small size of the colonies, exceeding a dozen individuals in a very few species (Baracchi et The phylogeny of the Stenogastrinae wasps is still under discus- al., 2009a), these organisms have been proposed as a sion and their systematic incomplete. In the present work we used geometric morphometrics, a technique based on a rigorous key group for understanding the origin of social evolu- statistical assessment of shape, to compare the forewings of fif- tion in insects (Yoshikawa et al., 1969; West-Eberhard, teen species of Stenogastrinae wasps belonging to four different 1978; Turillazzi, 1991). Several of their features, like genera to ascertain whether this approach may be used as a reli- the elaborate larval rearing (Turillazzi, 1989), the long able method in the study of the taxonomy of the group. The re- larval developmental time (Field et al., 2000) and the sults show that the wing vein junctions can be diagnostic for both genus and species identification. For the first time in this poor quality of nest material (Hansell, 1987) are con- subfamily, we propose a phylogenetic classification of the spe- sidered as important determinants in shaping their cies based on wing morphology that largely agrees with the peculiar social structure. The most generally accepted cladistic data available at genus level and reflects the differ- phylogenetic hypothesis (Carpenter, 1982, 1988; Car- ences among species in terms of nesting material and architec- penter and Starr, 2000) collocates them as the sister- ture of their nest. group of the Polistinae + Vespinae, rendering the so- cial wasps as an entirely monophyletic group. Howev- er, a recent study by Hines et al. (2007), comparing Contents mitochondrial and nuclear DNA, indicated a clade Introduction .................................................................................... 191 composed of these three subfamilies as non-monophy- Material and methods ................................................................... 192 letic and implied a separate route to eusociality for the Samples collection .................................................................. 192 Stenogastrinae. The noticeable discrepancy between Geometric morphometrics analyses ................................... 193 these two contrasting hypotheses underlines the need Results .............................................................................................. 193 for further studies in order to clarify the taxonomy of Geometric morphometrics analyses ................................... 193 Discussion ....................................................................................... 195 the Stenogastrinae wasps and to better understand Acknowledgements ....................................................................... 197 their systematic at the subfamily, genera and species References ....................................................................................... 197 levels. Data concerning the phylogenetic relationships among species within the group are still missing ex- cept for the ones recently obtained for a few species Introduction belonging to the Liostenogaster, Eustenogaster and Parischnogaster genera through chemical taxonomy The Stenogastrinae represent a sub-family of social (Baracchi et al., 2009b). Taxonomists usually identify wasps belonging to the family Vespidae (Carpenter, species using operational methods, based on phenotyp- 1982; Hines et al., 2007). Stenogastrinae are endemic ic morphological evidence, on molecular (phylogenet- to the Indo-Malaysian area and comprise 57 described ics) and on chemical characters (Agapow et al., 2004; species belonging to seven genera among which Eus- Haverty et al., 2005; Dapporto, 2007; Hines et al., 2007; tenogaster, Liostenogaster and Parischnogaster show Baracchi et al., 2009b). Furthermore, in the Stenogas- the highest number of taxa described so far (Carpen- trinae, characteristics of the nest architecture have also ter, 2001). Due to the primitive social organization and been used to discriminate between morphologically CtZ80-03 (3) Baracchi.indd 191 03-08-11 09:29 192 Baracchi et al. – Wing morphology in Stenogastrinae Fig. 1. Calculated landmark points posi- tioned at vein junctions, used in fore- wing geometric morphometrics analyses of Stenogastrinae species. similar species (Sakagami and Yoshikawa, 1968). whether wing morphology of Stenogastrinae shows Classical morphometric methods (Ruttner, 1988), any correlation with their phylogeny at the genus level. based on multiple measurement of many individuals Finally, we provide, for the first time in the subfamily, (Alpatov, 1929) evolved in geometric morphometrics a dendrogram showing a phylogenetic classification of (Bookstein, 1991). This technique is based on a rigor- the species based on wing morphology. ous statistical theory of shape (Kendall et al., 1999) that allows quantitative multivariate analyses on struc- ture shapes (Rohlf and Marcus, 1993; Adams et al., Material and methods 2004). While some authors suggest that the use of geo- metric morphometrics data for reconstruction of phy- Samples collection logenies could be problematic (MacLeod and Forey, 2002), there is also mounting evidence supporting the Forewings of the following Malaysian Stenogastrinae use of this technique in insect systematics studies. Ac- species were collected in the field and used in this tually, the geometric morphometrics applied to vein study: Eustenogaster calyptodoma (Sakagami and junctions of the wings has already been successfully Yoshikawa, 1968), E. micans (de Saussure, 1852), E. used to discriminate honeybee subspecies (Tofilski, fraterna (Bingham, 1897), Liostenogaster campanu- 2008), Sphex species (Tuzun, 2009), and some species lae (Turillazzi, 1999), L. flavolineata (Cameron, 1902), of Syrphidae (Francuski et al., 2009). L. nitidipennis (de Saussure, 1952), L. pardii (Turillazzi In the present work geometric morphometrics was and Carfì, 1996), L. topographica (Turillazzi, 1999), applied to forewings of 15 species belonging to four L. vechti (Turillazzi, 1988), Metischnogaster drewseni genera of Stenogastrinae wasps to assess whether this (de Saussure, 1857), Parischnogaster alternata (Sak- approach may contribute to taxonomic studies and agami, 1969), P. mellyi (de Saussure, 1852), P. jacob- Fig. 2. Graphical representation of the first (x-axis) and of the second (y-axis) Relative Warp (RW) of the forewing vein junctions analysis. Open penta- gons, E. calyptodoma; black stars, E. micans; open rhombus, E. fraterna; tri- ad, L. campanulae; black triangles, L. flavolineata; open triangles, L. ni- tidipennis; open circles, L. pardii; black rhombus, L. topographica; black rectan- gles, L. vechti; open squares, M. drews- eni; black circles, P. alternata; black squares, P. mellyi; X-shaped, P. jacob- soni; sun-shaped, Parischnogaster sp., open stars, P. striatula. CtZ80-03 (3) Baracchi.indd 192 03-08-11 09:29 Contributions to Zoology, 80 (3) – 2011 193 soni (du Buysson, 1913), Parischnogaster sp., P. striat- their species group. We provided the average land- ula (du Buysson, 1905). Parischnogaster sp. is an un- marks configuration because it can be used as a refer- described species but it is quite common in Malaya ence for future studies (Francoy et al., 2008). Moreo- peninsula (pers. obs. DB) and it belongs to the Par- ver, using the shape residual from GPA, the Partial ischnogaster genus characterized by a very narrow Warps, that are sets of variables containing shape in- petiole of the gaster and distinctly longer than wide, formation, were calculated. Applying the PCA princi- clypeus widely separated from the eye and posterior ple to Partial Warps, we obtained Relative Warps ocelli separated by more than one ocellus diameter. Its (RW) that could be used as variables in stepwise dis- nest is similar to that of P. jacobsoni but it is often criminant analysis (DA). RW can be visualized by spread on a flat surface like a leaf (with each cell sepa- thin-plate-spline deformation grids, which allow a rate from the others) and equipped with an Ant-guard. visual comparison of shape differences. GPA, Partial All the samples (ten females for each species, except for and Relative Warps calculations and thin-plate-spline M. drewseni for which only 9 females were used; each visualization were carried out using TPSRELW 1.45 specimen from a different colony) were collected in Pa- (Rohlf, 2007). Since the number of RW was high, only hang State in peninsular Malaysia during February- RW explaining more than 1% of variance were used in March 2008 and killed by freezing soon after capture. DA. The significance of Wilks’ lambda and the per- As nest architecture is important for species identifica- centage of correct assignments were used to estimate tion, we collected the specimens directly on the comb. the validity of the discriminant functions. Moreover, Since male wasps disperse soon after emergence, we we performed a complete cross-validation test using focused on females. Some specimens were
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