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Identification of Cryptic Species and Morphotypes in Male Euglossa: Morphometric Analysis of Forewings (Hymenoptera: Euglossini) J

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Identification of Cryptic Species and Morphotypes in Male Euglossa: Morphometric Analysis of Forewings (Hymenoptera: Euglossini) J Identification of cryptic species and morphotypes in male Euglossa: morphometric analysis of forewings (Hymenoptera: Euglossini) J. J. G. Quezada-Euán, H. D. Sheets, E. de Luna, T. Eltz To cite this version: J. J. G. Quezada-Euán, H. D. Sheets, E. de Luna, T. Eltz. Identification of cryptic species and morpho- types in male Euglossa: morphometric analysis of forewings (Hymenoptera: Euglossini). Apidologie, Springer Verlag, 2015, 46 (6), pp.787-795. 10.1007/s13592-015-0369-7. hal-01319634 HAL Id: hal-01319634 https://hal.archives-ouvertes.fr/hal-01319634 Submitted on 21 May 2016 HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. Apidologie (2015) 46:787–795 Original article * INRA, DIB and Springer-Verlag France, 2015 DOI: 10.1007/s13592-015-0369-7 Identification of cryptic species and morphotypes in male Euglossa : morphometric analysis of forewings (Hymenoptera: Euglossini) 1 2 3 4 J. J. G. QUEZADA-EUÁN , H. D. SHEETS , E. DE LUNA , T. ELTZ 1Departamento de Apicultura Tropical- Campus Ciencias Biológicas y Agropecuarias, Universidad Autónoma de Yucatán, Mérida, Mexico 2Canisius College, Buffalo, NY, USA 3Biodiversidad y Sistemática, INECOL, Xalapa, Mexico 4Department of Animal Ecology, Evolution and Biodiversity, Ruhr-Universität, Bochum, Germany Received 3 February 2015 – Revised 10 April 2015 – Accepted 17 April 2015 Abstract – Males of sibling orchid bees Euglossa viridissima and Euglossa dilemma are morphologically cryptic, except for the number and shape of mandibular teeth. An alternative morph of E. viridissima has a third tooth similar to males of E. dilemma . We used this model system to evaluate the potential of wing morphometrics for the resolution of these groups. We found differences in the size characters of forewings of E. viridissima and E. dilemma albeit with substantial overlapping amongst them. However, geometric morphometrics of forewing vein intersections separated both species and, to a lesser extent, morphotypes. A discriminant analysis of the shape of the radial cell showed separation between all three groups, too, albeit with higher misclassification between E. viridissima and E. dilemma . We show that sibling cryptic species and morphotypes can be identified by geometric morphometrics, supporting its application with other methods as powerful aids to infrageneric taxonomy in bees. orchid bee / cryptic species / morphotypes / geometric morphometrics / forewing 1. INTRODUCTION The taxonomy of Euglossini is mostly based on the analysis of male traits (Roubik and Hanson Euglossines, commonly called orchid bees 2004). Detailed morphological revisions of male (Hymenoptera, Apidae, Euglossini) are neotropi- Euglossini have resulted in the discovery of sev- cal, iridescent colored insects, well known for the eral sibling and cryptic species (Oliveira and males’ behavior of collecting volatiles from or- Nemésio 2003; Ramírez 2006; Rasmussen and chids and other sources (Dressler 1982). Euglossa Skov 2006; Nemésio and Bembé 2008; Letreille is the largest genus of orchid bees with Hinojosa-Díaz et al. 2011, 2012; Nemésio and over 100 described species (Roubik and Hanson Engel 2012). These types of revisions can make 2004). use of the additional information obtained from morphometric multivariate methods and molecu- lar plus biochemical tools (Eltz et al. 2011; Electronic supplementary material The online version of this article (doi:10.1007/s13592-015-0369-7) contains Pokorny et al. 2014). supplementary material, which is available to authorized Insect wing morphometrics has been shown to users. be a quick and inexpensive method that provides Corresponding author: J. Quezada-Euán, reliable resolution on the specific and infra- [email protected] specific levels of various taxa (Ruttner 1988;Haas Manuscript editor: Klaus Hartfelder and Tolley 1998; Klingenberg et al. 2001; 788 J.J.G. Quezada-Euán et al. Dujardin et al. 2003). Recently, geometric mor- centrally (Eltz et al. 2011). Females of both spe- phometric approaches have shown higher statisti- cies are superficially identical and can only be cal power compared to traditional morphometrics separated by molecular markers (Eltz et al. 2011; for species separation (Bookstein 1991;Zelditch May-Itzá et al. 2014). Male 2D and 3D et al. 2004; Francoy et al. 2011; Zinetti et al. E. viridissima are chemically and genetically in- 2013). By eliminating the effect of size, statistical distinguishable (Eltz et al. 2011; Pokorny et al. comparison of landmarks used in geometric mor- 2014; Ramírez et al., unpublished data). More- phometrics can reveal differences of actual shapes over, both male types emerge from clutches of or forms with greater power of discrimination the same mother (T. Eltz and Quezada-Euán, un- (Mitteroecker and Gunz 2009). In bees, geometric published data), confirming that they are in fact morphometrics of the forewing has been capable individuals of the same species. of discriminating operational taxonomic units In this study, we evaluated the power of wing (Aytekin et al. 2007; Francoy et al. 2008, 2011; morphometrics for identifying cryptic species and Kandemir et al. 2011). Importantly, such identifi- species morphotypes in bees using as a model the cation seems to conform to those based on molec- E. viridissima–E.dilemma species complex. Both ular markers (Francoy et al. 2012; Oleksa and traditional morphometrics based on wing size (tra- Tofilski 2015). However, in bees, the use of geo- ditional morphometrics) and geometric morpho- metric morphometrics for the discrimination of metrics of shape were included in the study. cryptic species has remained uninvestigated, in spite of its promising use in other insect taxa 2. MATERIALS AND METHODS (Baylac et al. 2003; Pizzo et al. 2006; Gurgel- Goncalves et al. 2011; Muñoz-Muñoz et al. 2.1. Material collection 2011; Mitrovski—Bogdanović et al. 2013). Over- all, the application of geometric morphometrics to Males of E. viridissima and E. dilemma were col- assess intraspecific morphological differentiation lected in 13 localities across the Yucatan Peninsula and intra-sexual plasticity is still a largely unex- during 2010 using synthetic benzyl benzoate (BB), plored field in bees (Danforth and Desjardins 1,8-cineole (C), eugenol (E), methyl salicylate (MS), 1999). p-dimethoxybenzene (PDMB), and methyl cinnamate In this respect, male orchid bees of sibling (TMC) as baiting substances (Pokorny et al. 2013). Euglossa viridissima Friese and Euglossa dilem- Individuals were kept in ethanol and stored at −20 °C ma Bembé and Eltz represent a unique model for awaiting further analyses. Males were separated into evaluating geometric morphometric methods in E. dilemma and E. viridissima and the two morphs in the identification of cryptic species and the latter in accordance to the number of mandibular morphotypes altogether. The E. viridissima–E. teeth and their position. For all statistical analyses, the dilemma species pair exists in sympatry in the three a priori groups consist of 44 2D and 23 3D Tehuantepec Isthmus and the Yucatan Peninsula specimens of E. viridissima , and 41 of E. dilemma of Mexico (Eltz et al. 2011). Morphometric were identified. methods based on measures of body size have failed to separate sibling E. viridissima and E. dilemma males which remain cryptic for sev- 2.2. Preparation of forewings eral characters traditionally used in bee morpho- metrics (Ramirez-Pech 2009). So far, males of From each specimen, the right forewing was dis- both species can only be separated phenotypically sected from the base of the radial vein with the help of by their number and shape of mandibular teeth. forceps and mounted between glass slides to keep Males of E. viridissima are commonly bidentate them flat. Wings were photographed using an (2D, >90 % of individuals) but occasionally OpticFilm 7200 scanner (Plustek) for diapositives at tridentate (3D) with the middle dentition near the 3600 dpi. We used 28×48 mm, 1.55-mm thickness distal one, whereas E. dilemma are always microscope slides to sandwich wings, which fitted tridentate with the middle dentition placed the slide scanner. Identification of cryptic species/morphotypes in male Euglossa 789 2.3. Analyses of size Table I. Distances and angles formed by venation land- marks used in the analyses of forewing size and shape. For the analyses of size, we used distances and angles formed by vein intersections. We first recorded Distances Landmark Landmark Landmark between vein 1 2 3 (angle 18 landmarks of the intersections of the veins on the intersections intersect) right forewing (Figure 1). Cartesian coordinates of points on the photograph of the forewing of each indi- 11314– vidual male were registered using the software tpsDig2 2113– version 2.12 (Rohlf 2008). After this, all configurations 31115– were aligned using Procrustes superimposition by 41216– means of the software CoordGen6 (Sheets 2006). 5816– Ten inter-landmark distances and seven angles be- 61012– tween vein intersections were then calculated using the 739– program TMorphGen (Sheets 2006) generating
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