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Medium Molecular Weight Polar Substances of the Cuticle As Tools In Accepted on 10 June 2009 Ó 2009 Blackwell Verlag GmbH J Zool Syst Evol Res doi: 10.1111/j.1439-0469.2009.00543.x 1Universita` degli Studi di Firenze, Dipartimento di Biologia Evoluzionistica ‘‘Leo Pardi’’, Via Romana, Firenze, Italy; 2Istituto Comprensivo Materna Elementare Media Convenevole da Prato via 1° Maggio, Prato, Italy; 3Institute of Biological Sciences, University of Malaya, Kuala Lumpur, Malaysia; 4Centro Interdipartimentale di Spettrometria di Massa dellÕUniversita‘ di Firenze, Viale G. Pieraccini, Firenze, Italy Medium molecular weight polar substances of the cuticle as tools in the study of the taxonomy, systematics and chemical ecology of tropical hover wasps (Hymenoptera: Stenogastrinae) David Baracchi1,Leonardo Dapporto2,Serafino Teseo1,Rosli Hashim3 and Stefano Turillazzi1,4 Abstract The Stenogastrinae wasps have been proposed as a key group for an understanding of social evolution in insects, but the phylogeny of the group is still under discussion. The use of chemical characters, in particular cuticular hydrocarbons, for insect taxonomy is relatively recent and only a few studies have been conducted on the cuticular polar substances. In this work, we ascertain, by the matrix-assisted laser desorption ionization- time of flight mass spectrometry technique, that different species of primitively eusocial hover wasps have different compositions of the epicuticular polar compounds ranging from 900 to 3600 Da. General linear model analysis and discriminant analysis showed that the average spectral profiles of this fraction can be diagnostic for identification of the species. Moreover, for the first time we show population diversification in the medium MW polar cuticular mixtures in insects. In conclusion, the results demonstrate that the chemical characters are consistent with the physical characters and the study support the importance of medium MW polar substances as powerful tools for systematics (chemosystematics) and chemical ecology (fertility signal and population characterization) in a primitively social insect taxon. Key words: Chemotaxonomy – epicuticular profiling – polar compounds – MALDI-TOF Introduction genera with the highest number of species (Liostenogaster van The Stenogastrinae wasps (Hymenoptera, Vespidae) are a taxon der Vecht, 1969, and Parischnogaster von Schulthess, 1914) of approximately 58 described species in seven genera inhabiting still lack a published revision. Classic taxonomic methods the forests of South-east Asia from South India to New Guinea based on differences in morphological character sets continue and from Vietnam to Indonesia (Carpenter and Starr 2000). The to be powerful tools for the discrimination of species. largest genera are Eustenogaster van der Vecht, 1969, Lioste- Nevertheless, in the Stenogastrinae, characters of nest archi- nogaster van der Vecht, 1969 and Parischnogaster von Schulth- tecture have also been used to discriminate among very similar ess, 1914 all distributed in the Oriental Region. These wasps, species (Sakagami and Yoshikawa 1968). The use of chemical owing to their small colonies and primitive social organization, characters for insect taxonomy is relatively recent. In partic- have been proposed as a key group for an understanding of ular, the cuticular hydrocarbon composition has been assessed origin and social evolution in insects (Yoshikawa et al. 1969; as a possible chemotaxonomic character in several taxa, such West-Eberhard 1978; Turillazzi and Pardi 1982; Turillazzi as mosquitoes (Carlson et al. 1997), beetles (Page et al. 1997), 1991), and several of their features are considered important in grasshoppers (Chapman et al. 1995), butterflies (Dapporto promoting or limiting their social evolution [e.g. poor quality of 2007) and, among social insects, ants (Vander Meer 1986), bees nest material, (Hansell 1987); elaborate larval rearing, (Turi- (Blum et al. 2000) and termites (Haverty et al. 2000, 2005). llazzi 1989); ensured fitness returns and long larval develop- Besides having genetic determinants, these compounds are also mental times, (Field et al. 2000)]. used in communication and thus may represent a interesting The phylogeny of the group is not completely clear: the link between DNA and ecological and behavioural traits prevailing view (Carpenter 1982, 1988) is that the three typical of each species (see Dapporto et al. 2004a,b,c for social subfamilies of social wasps form a monophyletic group with wasps). However, hydrocarbons are not the exclusive compo- the hover wasps as a sister group of the other two social vespid nents of the superficial cuticular layer of insects and more subfamilies Polistinae and Vespinae. However, a recent study precisely of wasps. In fact, mass spectrometry analysis of by Hines et al. (2007), comparing mitochondrial and nuclear cuticular methanol extracts from Polistes paper wasps showed DNA, indicated a clade composed of these three subfamilies as a complex pattern of polar substances (Dapporto et al. 2008), non-monophyletic and implied a separate route to eusociality including substances similar in part to those found in the for the Stenogastrinae. Hence, the systematics of the group is medium molecular weight fraction of venoms of various not yet complete: 58 species in seven genera are currently species (Turillazzi et al. 2007). Moreover, it has been also recognized (Carpenter 2001). The genus Eustenogaster van der demonstrated that cuticular hydrocarbons may derive directly Vecht was recently revised (Saito and Kojima 2007), but the from prey and thus may be strongly dependent on environ- mental factors (Liang and Silverman 2000). Conversely, venom and cuticular peptides are likely produced directly by Corresponding author: David Baracchi ([email protected]) protein synthesis and may represent a more stable indication AuthorsÕ email addresses: Leonardo Dapporto ([email protected]), of phylogenetic relationships. Serafino Teseo ([email protected]), Rosli Hashim (roslihashim@ The aims of the present study were to ascertain whether um.edu.my) and Stefano Turillazzi ([email protected]) medium molecular weight polar compounds are present on the J Zool Syst Evol Res (2010) 48(2), 109–114 110 Baracchi, Dapporto, Teseo, Hashim and Turillazzi cuticle of hover wasps and, if so, whether they have a species- samples, was analysed using an automatic procedure available on the specific composition and could possibly be used to discrimi- MALDI mass spectrometer to standardize the results. nate between different taxa. We used the matrix-assisted laser Methanol extracts from each individual wasp were analysed with a MALDI Ultraflex TOF⁄TOF (Bruker Daltonics, Bremen, Germany). desorption ionization-time of flight mass spectrometry (MAL- The instrument was operated in positive ion reflector mode. The DI-TOF MS) technique to determine the reference spectra accelerating voltage and the Ion Source 2 were set to 25.0 and 21.9 kV, (profiling) of the cuticular layer of some species belonging to respectively, and the delay time was 20 ns; 800 shots were automat- three genera of Stenogastrinae. Peptide mapping and spectra ically accumulated for each spectrum. External calibration was profiling using MALDI-TOF MS analysis and other MS performed using the Bruker Standard Peptide Calibration kit (Bruker techniques have already been used to demonstrate differences Daltonics, Bremen, Germany). flex analysis between individuals belonging to the same species (Jakubowski We also performed an internal calibration using the software by Bruker Daltonics, Bremer, Germany, as we had the exact et al. 2005) or to establish the phylogenetic relationships mass of two peptides (1854 and 1909 Da) identified in the venom of the among various species belonging to the same taxonomic group paper wasp Polistes dominulus. Calibrated spectra were imported into (Serrao Wermelinger et al. 2005). In social Vespidae, MS the clinprotoolsÔ (cpt) software (Bruker Daltonics, Bremen, techniques have been employed for the study of medium MW Germany) and processed with a procedure suggested by the software venom fractions from some species of social wasps (Turillazzi manual and similar to that used by Zhang et al. (2004) for profiling et al. 2007) and for the characterization of the main allergens potential biomarkers in the plasma of asthma patients and to that of a previous study in our laboratory on cuticular and venom peptides of the venoms of various species (Kolarich et al. 2005). (Turillazzi et al. 2006). The program calculates the areas of the most Recently, our group used GC–MS techniques to characterize important peaks that may account for statistical differences between volatile hydrocarbons and other apolar compounds of the various species. This data set was used for statistical analyses using spss venom of Polistes wasps (Bruschini et al. 2006a,b, 2007). The 13 for Windows (SSPS, Inc., Chicago, IL, USA). MALDI technique requires very low amounts of substance, allowing us to perform analyses, in a very short time, of the Statistical analyses epicuticular polar substances of single individuals belonging to each examined species. In this paper, we report the results We first determined the percentage of each peak area with respect to obtained from analyses of epicuticular polar odours ranging the total of all peaks and then we log transformed it. We performed a principal components analysis (PCA) to reduce the number of from 900 to 3000 Da collected from 11 tropical hover wasp variables subsequently used in general linear model (GLM) analysis species belonging
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