Hymenoptera: Megachilidae) in a Monte Desert Reserve in Mid-Western Argentina Nydia Vitale, Diego P
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Ecology and nesting biology of the wood-boring bee Trichothurgus laticeps (Hymenoptera: Megachilidae) in a Monte desert reserve in mid-western Argentina Nydia Vitale, Diego P. Vázquez To cite this version: Nydia Vitale, Diego P. Vázquez. Ecology and nesting biology of the wood-boring bee Trichothurgus laticeps (Hymenoptera: Megachilidae) in a Monte desert reserve in mid-western Argentina. Apidologie, Springer Verlag, 2017, 48 (1), pp.31-40. 10.1007/s13592-016-0446-6. hal-01532475 HAL Id: hal-01532475 https://hal.archives-ouvertes.fr/hal-01532475 Submitted on 2 Jun 2017 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 (2017) 48:31–40 Original article * INRA, DIB and Springer-Verlag France, 2016 DOI: 10.1007/s13592-016-0446-6 Ecology and nesting biology of the wood-boring bee Trichothurgus laticeps (Hymenoptera: Megachilidae) in a Monte desert reserve in mid-western Argentina 1 1,2 Nydia VITALE , Diego P. VÁZQUEZ 1Instituto Argentino de Investigaciones de las Zonas Áridas, CONICET, 5500, Mendoza, Argentina 2Facultad de Ciencias Exactas y Naturales, Universidad Nacional de Cuyo, M5502JMA, Mendoza, Argentina Received 14 January 2016 – Revised 17 March 2016 – Accepted 31 March 2016 Abstract – Studying bee nests can enlighten our understanding of feeding specialization and phylogenetic relationships of bees. We studied the nesting and feeding habits of Trichothurgus laticeps in the Monte desert ecosystem. Our results show that T. laticeps is attracted to pre-existing cavities in wood (trap nests), which were further excavated for nest construction; nest cells are unlined and provisioned almost exclusively with pollen of the cactus Opuntia sulphurea . The species overwinters inside a hard cocoon. Our results reinforce the hypothesis previously proposed by other authors that the unworked pollen mass and oligolecty are plesiomorphic characters in the Megachilidae, and that the wood-boring behavior for nest construction indicates convergent evolution with the Xylocopinae. We also propose that the cocoon structure might represent an evolutionary novelty for the Lithurginae. host-plant specialization / Lithurginae / nesting behavior and structure / phylogenetic relationships / solitary bees 1. INTRODUCTION Megachilinae (Danforth et al. 2013; Hedtke et al. 2013). It is accepted that Fideliinae is the sister The study of bee nests, including their architec- clade to the remainder of the family (Danforth et ture, the structure of brood cells, the type of provi- al. 2013;Hedtkeetal.2013;Litmanetal.2011; sions, and larval behavior, may enhance our under- Michener 2007). While morphological studies standing of the ecology and evolution of bees (Gonzalez et al. 2012) indicate that the (Michener 2007). The Megachilidae is an impos- Pararhophitinae are more basal than the ing bee family, with over 4000 described species Lithurginae, molecular studies are less conclusive, (Ascher and J. Pickering 2010). Phylogenetic stud- given that both groups exhibit several ies suggest this family is monophyletic (Danforth plesiomorphic characters (Hedtke et al. 2013; et al. 2013;Hedtkeetal.2013; Litman et al. 2011; Litman et al. 2011). Roig-Alsina et al. 1993) with four subfamilies: Studying bee nests can also illuminate our un- Fideliinae, Lithurginae, Pararhophitinae, and derstanding of feeding specialization in bees. Although a number of bee species have been iden- tified as purported pollen specialists (e.g., Neff Electronic supplementary material The online version of 2003;NeffandSimpson1992), in many cases, this this article (doi:10.1007/s13592-016-0446-6) contains supplementary material, which is available to authorized conjecture rests primarily on visitation observations users. or scopal pollen loads, which usually represent a single foraging trip (Sipes and Tepedino 2005). In Corresponding author: N. Vitale, [email protected] contrast, pollen provisions in bee nests offer copi- Manuscript editor: Klaus Hartfelder ous information on the foraging habits of female 32 N. Vitale and D.P. Vázquez bees, integrating their resource use in space and Our study focuses on Trichothurgus laticeps time. Several studies have so far attempted to eval- Friese, 1906, a species occurring in xeric regions uate pollen specialization in bees using pollen pro- of Argentina. We studied the foraging and nesting visions in brood cells (Bosch et al. 1993;Buschini biology of T. laticeps , including cell provisions, et al. 2009; Dórea et al. 2013; Gonçalves et al. 2012; cocoon structure, and nest architecture. Rust et al. 2004; Schlindwein et al. 2009). The available information on nesting and feed- 2. MATERIAL AND METHODS ing habits is extremely limited for some groups in the Megachilidae, which hinders our understand- 2.1. Study sites and field methods ing of their phylogenetic relationships. This is the case for the Lithurginae, for which the information Data used in this study come from field studies on nesting habits is particularly limited. This sub- conducted between 2006 and 2012 in the Monte family includes four genera: Lithurgus , Desert of Villavicencio Nature Reserve, Mendoza, Lithurgopsis , Microthurge , and Trichothurgus . Argentina (Chacoff et al. 2012;Dorado2011; There is information about the nesting biology of Vázquez et al. 2009, 2012; Table I). Because data come the former three genera from several studies, with from several projects running simultaneously at the at least 12 studies on Lithurgus and Lithurgopsis same sites, there are minor differences in the sampling species (Rozen 2013; Rozen and Hall 2014), one methods used in different years. In 2006 and 2007, four study on one Microthurge species (M. corumbae ; sample sites were located near a dry temporary stream at Garófalo et al. 1992), and additional notes on the ca. 1240 m above sea level. In 2008, data were collected occurrence of other species in South America at 18 sites located at different elevations above sea level (Ascher and J. Pickering 2010; Moure and Melo (range 1078–1450 m), post fire age (range 1–50+ 2012). Most species in these three genera excavate years), and vegetation types. In 2009–2011, data were their burrows in wood and construct their nests with collected at six sites located at 1115–1750 m above sea unlined brood cells (Rozen 2013); trophically, they level. Additional sites were included in 2012, range from oligolectic, feeding on few plant species expanding the elevational range of the studies. (Díaz and Cocucci 2003), to polylectic, feeding on Data on pollinator visitation to flowers came from a many plant species of multiple families (Güler and study conducted between 2006 and 2011 (Chacoff et al. Sorkun 2007). The information available for 2012 and additional unpublished results). Pollinator Trichothurgus is comparatively rather limited, visits to flowers were recorded in the spring and early coming from anecdotal nesting observations of summer (September to December or January) in six T. dubius (Claude-Joseph 1926;Rozen1973)and consecutive flowering seasons. We recorded flower vis- astudyofT. bolithophilus in which this species iting insects on plant species in weekly surveys, sam- was observed nesting in dry horse dung (Sarzetti pling the whole community as comprehensively as pos- et al. 2012). Although Trichothurgus species were sible (see Chacoff et al. 2012 for further details on believed to feed primarily on pollen from cactus sampling methods). (Michener 2007), T. bolithophilus was recorded Bees were studied using wooden trap nests. Each feeding primarily on pollen from the Asteraceae study site was a 200 × 100 m rectangular area with six and Amaranthaceae (Sarzetti et al. 2012). Thus, it sampling points separated by 100 m. Each sample point is unclear whether the anecdotic records of the nest consisted of two poles separated by 10 m. Each pole building and provisioning habits of T. dubius and contained 24 traps of three different sizes: eight traps of T. bolithophilus are representative of the diversity 14 cm hole length and 0.5 cm of hole diameter, eight of nesting habits of Trichothurgus , and whether the traps measuring 14 cm × 0.8 cm, and eight traps mea- above observations indicate a departure from the suring 28 × 1.1 cm; the smaller traps were placed at wood nesting habits of Lithurgus and Microthurge . 0.70 m from the ground, the longer at 0.40 m. All study Answering these questions should improve our sites were visited weekly to conduct plant abundance understanding of the evolution of the nesting habits surveys and to collect nests in the spring and summer, of lithurgine bees and of the phylogenetic relation- the flowering and bee activity season in the study area. ships within this group and with other bee groups. We used all data sets to describe the natural history of Ecology and nesting biology of T. laticeps 33 Table I. Resume of study aims and data use for the T. laticeps nest sites in a Monte desert reserve. Aim Year Number of sites Number of nests built Natural history (whole study) 2007–2012 21 221 Phenology 2009 1 11 Visits to flowers (census) 2006–2011 2 – Temporal co- 2007–2012 2 42 occurrence Spatial co- 2008 18 21 occurrence Larval pollen 2008–2009 11 80 provision Parasite attack 2012 1 1 T. laticeps collected between 2008 and 2012 in the sites with the species level function of R’s package described above. For the analysis of plant/bee spatial bipartite. The index d measures the deviation co-occurrence, we used the 2008 data. To study inter- from randomly distributed interactions, and varies annual variation in nesting habits, we used the 2008– between 0 (extreme generalization) and 1 (extreme 2011 data for the two sites for which we had uninter- specialization).