Hymenoptera, Apidae) in Argentina
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Agricultural and Forest Entomology (2017), DOI: 10.1111/afe.12207 Nesting ecology and floral resource of Xylocopa augusti Lepeletier de Saint Fargeau (Hymenoptera, Apidae) in Argentina ∗† †‡ ∗† ∗† Mariano Lucia , María C. Telleria , Pablo J. Ramello and Alberto H. Abrahamovich ∗División Entomología, Museo de La Plata, Universidad Nacional de La Plata, Edificio Anexo Museo, Unidades de Investigación FCNyM, 122 y60, 1900FWA La Plata, Argentina, †CONICET, Consejo Nacional de Investigaciones Científicas y Técnicas, La Plata, Argentina and ‡Laboratorio de Sistemática y Biología Evolutiva - Museo de La Plata, Universidad Nacional de La Plata, Paseo del Bosque s/n, 1900FWA, La Plata, Argentina Abstract 1 A total of 33 nests of Xylocopa augusti was studied during two consecutive seasons. 2 Nesting behaviour and floral resources used by the large carpenter bee X. augusti Lepeletier de Saint Fargeau were studied during the brood production season in an urban area in Argentina. 3 Biological information about nesting aspects inside and outside the nest was consid- ered, paying particular attention to year-long activity, foraging flights throughout the day for nectar and pollen collection, nectar dehydration, oviposition, and pollen pref- erence. 4 In the study area, X. augusti shows an univoltine life cycle, with a peak of nesting between October and December, which coincides with the greatest blooming period of the surrounding flora. 5 From 36 analyzed larval provision samples, 18 pollen types were identified, most of them belonging to ornamental trees or shrubs. Pollen from Eucalyptus- Myrceugenia glaucescens (Cambess.) D. Legrand and Kausel (Myrtaceae), Solanum sp.-Cyphomandra betacea (Cav.) Sendtn. (Solanaceae) and Erythrina crista-galli L. (Fabaceae) was dominant. 6 The ability to obtain pollen from poricidal anthers such as those of Solanum indicate the potential of X. augusti to be an excellent managed pollinator and a good candidate for pollinating Solanaceae, such as Solanum lycopersicum ‘tomato’ and Solanum melongena ‘eggplant’, which are economically important crops in this region. Keywords Argentina, floral resources, large carpenter bees, nesting behaviour. Introduction bees (Osmia spp.) are managed as commercial pollinators and used in commercial agriculture (Thorp, 2003; Torchio, 2003; The need for insect pollination in modern agriculture is a vital Velthuis & van Doorn, 2006; Pitts-Singer & Cane, 2011). yet expensive factor in crop production (Abrol, 2012). Apis The introduction of non-native managed pollinators for crop mellifera L. (Hymenoptera: Apidae) is usually suggested as the pollination and honey production can impact on native bee most important managed pollinator in commercial agriculture species through competition for resources or direct interaction worldwide (Free, 1993; Kearns et al., 1998). However, it has been and the introduction of pests and diseases (Goulson, 2003; long demonstrated that honeybees are not efficient pollinators for Plischuk et al., 2009; Schmid-Hempel et al., 2014; Alvarez et al., all crops (O’Toole, 1997; Garibaldi et al., 2013); for example, 2015). Among the wild bees that were tested in different crops for those plants that require vibratile or buzz pollination (i.e. the and considered as potentially manageable pollinators are species extraction of pollen from poricidal anthers using vibrations of of the genus Xylocopa (Hogendoorn et al., 2000; Freitas & the flight muscles) (Buchmann, 1983), some species of wild bees Oliveira Filho, 2001; Buchmann, 2004). Commonly known as are more effective. However, only a few species of bumblebees large carpenter bees, species of the genus Xylocopa Latreille (Bombus spp.), leaf-cutting bees (Megachile spp.) and mason (Apidae: Xylocopini) comprise conspicuous, large, hairy bees represented by 31 subgenera worldwide, with approximately 470 Correspondence: Mariano Lucia. Tel.: +54 9 221 4228451; e-mail: species being described, most of them occurring in tropical and [email protected] subtropical areas of the world (Lucia et al., 2015). Species of © 2017 The Royal Entomological Society 2 M. Lucia et al. Xylocopa commonly nest in solid wood, including structural The climate of the area is temperate, with an average tempera- timbers, or in dead stalks of plants, although some of them ture of 22 ∘C in the warmest month and 8 ∘C in the coldest; there nest in the ground. Their social behaviour varies across species, is no dry season and the average annual precipitation ranges from from solitary to semisocial or primitively eusocial nests where 800 to 1000 mm (Cicchino et al., 2003). the oldest female (mother or sister) feeds both young females and males via trophallaxis (Lucia et al., 2015). Compared with other non-Apis bees, carpenter bees are potentially more versatile Nest observations for agricultural pollination services because of their polylectic habits, the ability to buzz pollinate, long activity seasons, Xylocopa augusti is a native, parasocial bee widely distributed in hibernation as an adult, foraging when temperatures reach several countries of southern South America and is recorded in acceptable levels and the acceptance of artificial substrates for several provinces in Argentina. It is more widespread in the east, nesting (Buchmann, 2004; Keasar, 2010). Around the world, ranging from Buenos Aires to Formosa province, at the northern some of these species are effective pollinators on diverse crops, edge of its range (Lucia et al., 2014). This species constructs including on passion fruit, sunflowers, squash, tomato and nests in the dead wood of tree trunks or branches of a wide eggplant, and they are already being used for this purpose in diversity of plant families. The nests are structurally branched, different countries (Gerling et al., 1989; Sihag, 1993a; Mardan, with an entrance hole leading to a system of tunnels (Lucia 1995; Hogendoorn et al., 2000; Aguiar-Menezes et al., 2002; et al., 2014) that may be continuously occupied and expanded Sadeh et al., 2007). for several generations. The observations were conducted in One of the first steps to developing a management system that active artificial nests found in trap-nests that had been previously uses a bee pollinator is to understand its biology (i.e. life cycle, deployed for the purpose of the present study. The trap-nests nesting behaviour and floral preferences). The nesting biology consisted of internodes of canes (Arundo donax, Poaceae) closed of several Xylocopa species has been studied in Brazil (Camillo at one end (by the node) and open at the other, with the size of the et al., 1986; Bernardino & Gaglianone, 2008; Marchi & Melo, entrance varying from 12.50 to 17.00 mm, and they were placed 2010), Thailand (Hongjamrassilp & Warrit, 2014), India (Sihag, horizontally on open shelves in the field. Observations were 1993b, 1993c; Raju & Rao, 2006) and Israel (Gerling et al., made between 07.00 and 17.00 h by one observer during two 1983), although no such studies exist for Argentinian species. nesting periods (from October to April); nesting observations In the present study, we observe and describe the nesting (nest activities) were made from October to early January; and behaviour and floral resources used by X. augusti in an urban information about activity, mortality and founded nests was area in northeastern Buenos Aires province, Argentina. The recorded from October to April of each season. A total of 287 h results obtained should provided useful information for use (17.220 min) of observations was recorded in the field during the when evaluating this species as potential manageable pollinator nesting seasons. The sequence and duration of the inside/outside and also promote the use of wild bees for crop pollination, activities of each nest were observed and recorded, including: (1) especially those crops of particular interest commonly cultivated inside the nest: (i) digging of brood cells; (ii) nectar dehydration: by small-scale farmers in the studied area, such as Capsicum b1: at nest entrance and b2: inside the nest; (iii) deposition of spp. ‘peppers’, Solanum lycopersicum ‘tomatoe’ and Solanum pollen; (iv) preparation of provision mass; (v) construction of melongena ‘eggplant’. cell partition; (vi) egg laying (oviposition); and (vii) inactivity; (2) outside the nest (foraging activities): (i) nectar collection and (ii) pollen collection. The nests were classified as ‘founded nest’, Materials and methods when the trap nest was occupied by a female for the first time; Study area ‘active nest’, when the female occupies the nest and carry out the nesting activities; and ‘dead nest’, when the founder female Field work was conducted at the Unidad de Vivero Forestal Uni- dies for some reason and the nest becomes inactive. A flashlight ∘ ′ ′′ ∘ ′ ′′ versidad Nacional de La Plata (34 54 39 S, 57 55 37 W, was used to record observations inside the nest and, aiming to 18 m a.s.l.) located in La Plata city, Argentina. The area determine whether the foraging trip was for nectar or pollen comprises a total of 4 ha with a variety of cultivated trees collection, the presence of pollen in the scopa was considered as accompanied by weeds and adventitious species. The dominant a pollen foraging flight and absence was considered as a nectar trees belong to diverse families such as Platanaceae (Platanus foraging flight. Records of temperature and precipitation in the occidentalis L.), Tiliaceae (Tilia sp.), Myrtaceae (Eucalyp- study area were obtained from the ‘Departamento de Sismología