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Thoracobombus from Mexico: a Description of the Male Species Thoracobombus from Mexico: a description of the male species-specific cephalic labial gland secretions Nicolas Brasero, Rémy Vandame, Philippe Sagot, Baptiste Martinet, Irena Valterová, Pierre Rasmont To cite this version: Nicolas Brasero, Rémy Vandame, Philippe Sagot, Baptiste Martinet, Irena Valterová, et al.. Tho- racobombus from Mexico: a description of the male species-specific cephalic labial gland secretions. Apidologie, Springer Verlag, 2019, 50 (2), pp.183-194. 10.1007/s13592-018-0629-4. hal-02495376 HAL Id: hal-02495376 https://hal.archives-ouvertes.fr/hal-02495376 Submitted on 2 Mar 2020 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 (2019) 50:183–194 Original article * INRA, DIB and Springer-Verlag France SAS, part of Springer Nature, 2019 DOI: 10.1007/s13592-018-0629-4 Thoracobombus from Mexico: a description of the male species-specific cephalic labial gland secretions 1 2 2 1 Nicolas BRASERO , Rémy VANDAME , Philippe SAGOT , Baptiste MARTINET , 3,4 1 Irena VALTEROVÁ , Pierre RASMONT 1Research Institute of Biosciences, Laboratory of Zoology, University of Mons, Place du Parc 20, 7000, Mons, Belgium 2Departamento Agricultura Sociedad y Ambiente, El Colegio de la Frontera Sur (ECOSUR), San Cristóbal de Las Casas, Chiapas, Mexico 3Institute of Organic Chemistry and Biochemistry, Academy of Sciences of the Czech Republic, Flemingovo nám 2, CZ- 166 10, Prague, Czech Republic 4Faculty of Tropical AgriSciences, Czech University of Life Sciences, Kamýcká 129, CZ-165 21, Prague, Czech Republic Received 14 May 2018 – Revised 5 November 2018 – Accepted 14 December 2018 Abstract – The male cephalic labial gland secretions of bumblebees are known to be species-specific semiochemicals. These secretions that are involved in bumblebee pre-mating recognition provide efficient diagnostic characters for species delimitation. The subgenus Thoracobombus is the largest group of bumble- bees and is found in the Palearctic, Nearctic, and Neotropical regions. Here, the cephalic labial gland secretion (CLGS) composition of six Mexican Thoracobombus bumblebee species are analyzed: Bombus diligens , B. medius , B. mexicanus , B. pensylvanicus , B. trinominatus , and B. weisi . Our results suggest the presence of two new potential species into the formerly recognized B. weisi as well as one new potential species in the taxon presently identified as B. pensylvanicus . Moreover, the male of B. pensylvanicus , known to congregate at nest sites awaiting the emergence of virgin queens, is characterized by low concentrations of the C16 component. This observation raises the possibility that courtship behavior as well as environmental con- straints could affect the role of the male bumblebees’ CLGS. Thoracobombus / Mexico / cephalic labial glands / courtship behavior / species-specific 1. INTRODUCTION species are much more common and diverse in temperate areas of Eurasia and North Amer- The family Apidae (Hymenoptera) contains ica than in Neotropical region, where six 18 tribes including Bombini. This tribe is com- subgenera and 42 species have been recorded posed of social bees of the genus Bombus (Abrahamovich and Diaz 2002). Indeed, com- (Michener 2007) and currently includes about pared to the Palearctic region, the Neotropical 260 species in 15 subgenera (Williams 1998; region appears to have a less investigated bum- Williams et al. 2008). The majority of these blebee fauna (Williams 1998). This difference in number of recorded species could be ex- Electronic supplementary material The online version of plained by (i) a low sampling efforts in tropical this article (https://doi.org/10.1007/s13592-018-0629-4) regions compared to temperate ones, compli- contains supplementary material, which is available to cated by the difficulties in collecting specimens authorized users. in these regions and therefore the presence of a Corresponding author: N. Brasero, large number of undiscovered species and (ii) [email protected] the low number of researchers sampling and Manuscript editor: Bernd Grünewald determining bumblebees. 184 N. Brasero et al. After the studies on western hemisphere bum- mainly aliphatic compounds and are synthesized blebees carried out by Franklin (1913) and Frison de novo (Luxová et al. 2003; Žáček et al. 2013). (1925), the most recent revisions are those of They are known to be species-specific semio- Milliron (1971, 1973a, b). Other regional studies chemicals (Ayasse and Jarau 2014;Calam1969; were carried out in Brazil (Eustáquio Santos Terzo et al. 2003). These secretions are involved Júnior et al. 2015; Françoso et al. 2016; Moure in bumblebee pre-mating recognition by attracting and Sakagami 1962), Mexico and Central Amer- conspecific virgin females (Ayasse and Jarau ica (Labougle 1990), Colombia (Lievano-Leon 2014) but additional analyses are needed to deter- et al. 1994), Argentina (Abrahamovich and Diaz mine which compounds are directly involved in 2001), and Peru (Rasmussen 2003). In Mexico, 19 sexual recognition. Moreover, they provide effi- species belonging to 5 subgenera were recorded cient diagnostic characters for species delimitation (Abrahamovich and Diaz 2002;Franklin1913; (Bergström et al. 1981; Brasero et al. 2015, 2017; Frison 1925; Labougle 1990;Michener2007; Lecocq et al. 2015; Rasmont et al. 2005; Svensson Milliron 1971, 1973a). and Bergström 1979; Terzo et al. 2003). Within The Thoracobombus subgenus is the largest bumblebees, cephalic labial gland secretions of 88 group of bumblebees and is found in the Pale- species are currently known (Brasero 2018). In arctic, Nearctic, and Neotropical regions Thoracobombus , as well as in other subgenera, (Williams 1998). However, despite its large dis- the majority of compounds are shared by a large tribution, it remains quite rare and localized in number of species. Moreover, as already shown in the New World (Cameron et al. 2011; Milliron other insect groups such as beetles (Symonds and 1973a). Within this subgenus, nine species have Elgar 2004), it does not seem to be a phylogenetic been registered in Mexico. Compared to most pattern in the pheromones of bumblebees. bumblebees living in temperate regions In this paper, we aim to characterize the CLGS (Laverty and Plowright 1985), some additional composition of six Mexican Thoracobombus social characteristics such as the development of bumblebee species in order to highlight diver- large colonies (Michener and LaBerge 1954; gences in their sexual recognition system: Taylor and Cameron 2003; Zucchi 1973), or B. diligens , B. medius , B. mexicanus , well-developed defense systems (Cameron B. pensylvanicus , B. trinominatus ,andB. weisi . et al. 1999;Dias1954) have been highlighted in some Neotropical Thoracobombus species. 2. MATERIAL AND METHODS Mexico is a highly diverse country with large deserts (Chihuahuan Desert), mountain ranges 2.1. Sampling (Sierra Madre Occidental, Sierra Madre Oriental, Trans-Mexican volcanic belt), and highlands Sixty-nine male Thoracobombus specimens (Altiplano) and is biogeographically split by the were collected in Mexico between 2015 and Tehuantepec Isthmus in the south. These charac- 2016. They belong to 6 species: Bombus diligens teristics are favorable to the diversification of (n =1),B. medius (n =9),B. mexicanus (n =2), animals such as birds, insects, and lizards (Bryson B. pensylvanicus (n =8), B. trinominatus (n = et al. 2012; Cognato et al. 2008; Rodríguez- 30), and B. weisi (n = 19) (Supplementary file 1, Gómez et al. 2013). However, in the majority of Figure 1). All specimens were identified using the case, while geographic isolation combined references keys (Labougle et al. 1985;Milliron with an interruption of gene flow between popu- 1973a, b; Williams et al. 2008) and compared to lations is recognized as one of the main speciation the reference collection of El Colegio de la drivers (Coyne and Orr 2004; Sobel et al. 2010), Frontera Sur (ECOSUR) in Mexico. Following other process of divergence involving the choice the protocol found in De Meulemeester et al. of the sexual partner also play an important role 2011, males were killed by freezing at − 20 °C (Panhuis et al. 2001; Vereecken et al. 2007). and their CLGS were extracted in n -hexane The male cephalic labial gland secretions (400 μl). Samples were stored at − 40 °C prior to (CLGS) are complex mixtures composed of the analyses. Chemical description of bumblebees from Mexico 185 Figure 1 Maps of sampling sites of B. medius , B. diligens , B. pensylvanicus , B. mexicanus , B. trinominatus , and B. weisi . 2.2. Chemical trait differentiation CLGS compounds were quantified using a gas chromatography-flame ionization detector CLGS compositions were determined by gas Shimadzu GC-2010 equipped with an SLB-5ms chromatography-mass spectrometry (GC/MS) non-polar capillary column (5% phenyl (methyl) using a Focus GC (Thermo Scientific) with a polysiloxane stationary phase; 30-m column non-polar DB-5 ms capillary column (5% phenyl length; 0.25-mm inner diameter; 0.25-μm film (methyl) polysiloxane stationary phase;
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