Nectar Production Dynamics and Daily Pattern of Pollinator Visits in Brazil
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Nectar production dynamics and daily pattern of pollinator visits in Brazil nut (Bertholletia excelsa Bonpl.) plantations in Central Amazon: implications for fruit production Marcelo C. Cavalcante, Leonardo Galetto, Marcia M. Maués, Alípio José S. Pacheco Filho, Isac Gabriel A. Bomfim, Breno M. Freitas To cite this version: Marcelo C. Cavalcante, Leonardo Galetto, Marcia M. Maués, Alípio José S. Pacheco Filho, Isac Gabriel A. Bomfim, et al.. Nectar production dynamics and daily pattern of pollinator visits inBrazil nut (Bertholletia excelsa Bonpl.) plantations in Central Amazon: implications for fruit production. Apidologie, Springer Verlag, 2018, 49 (4), pp.505-516. 10.1007/s13592-018-0578-y. hal-02155119 HAL Id: hal-02155119 https://hal.archives-ouvertes.fr/hal-02155119 Submitted on 13 Jun 2019 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 (2018) 49:505–516 Review article * INRA, DIB and Springer-Verlag France SAS, part of Springer Nature, 2018 DOI: 10.1007/s13592-018-0578-y Nectar production dynamics and daily pattern of pollinator visits in Brazil nut (Bertholletia excelsa Bonpl.) plantations in Central Amazon: implications for fruit production 1,2 3 4 Marcelo C. CAVA LC AN TE , Leonardo GALETTO , Marcia M. MAUÉS , 1 1 1 Alípio José S. PACHECO FILHO , Isac Gabriel A. BOMFIM , Breno M. FREITAS 1Departamento de Zootecnia, Universidade Federal do Ceará, Avenida Mister Hull 2977, Campus do Pici, Fortaleza, CE CEP 60021-970, Brazil 2Universidade Federal Rural do Pernambuco, Unidade Acadêmica de Serra Talhada, Av. Gregório Ferraz Nogueira, s/n., Serra Talhada, PE CEP 56909-535, Brazil 3Departamento de Diversidad Biológica y Ecología e Instituto Multidisciplinário de Biologia Vegetal (UNC-CONICET), Universidad Nacional de Córdoba, Casilla de Correo 495, 5000, Córdoba, Argentina 4Laboratório de Entomologia, Embrapa Amazônia Oriental, Trav. Dr. Enéas Pinheiro s/n, Belém, PA 66095-903, Brazil Received 23 September 2017 – Revised 8 March 2018 – Accepted 26 April 2018 Abstract – We investigated composition and secretion patterns of nectar in the Brazil nut tree (Bertholletia excelsa ) and visitation patterns and glossa length of the main flower visitors along the anthesis, aiming to understand the implications for pollination and fruit production. Nectar sugar composition was dominated by sucrose and nectar secretion was continuous until 15:30 h, although flowers secreted, respectively, almost 50 and 80% of the total nectar volume and solutes in the hours immediately following flower opening, which coincides with peak flower visitation by bees. We observed a total of 19 bee species visiting the flowers to collect nectar throughout the day that can be considered pollinators. The three most abundant bee species were Xylocopa frontalis , Eufriesea flaviventris ,and Eulaema mocsaryi that accounted for about 90% of the visits. In open flowers, nectar was generally scarce, encouraging bees to move among trees, and likely increasing xenogamous pollen transfer in natural habitats. However, in the large-scale Brazil nut tree plantation studied here, where genetically identical (clone) individuals are planted together in high densities, even where bees move between trees, they seem to promote functional geitonogamy, determining pollen limitation. floral reward / foraging strategies / bee-plant interaction / pollination / reproductive success 1. INTRODUCTION (fructose, glucose, and sucrose) prevail in the total solutes (Stiles and Freeman 1993; Proctor et al. Nectar is one of the most important resources 1996). The production of nectar represents a costly offered by angiosperms to potential pollinators and, investment of plant resources, demanding a bal- although it may contain an ample variety of chem- anced trade-off between providing sufficient quan- ical components, three ordinary carbohydrates tity and quality to attract pollinators and maintain their interest in visiting other flowers, ensuring Electronic supplementary material The online version of cross-pollination, but at the lowest possible cost this article (https://doi.org/10.1007/s13592-018-0578-y) to the plant (Devlin and Stephenson 1985). Ac- contains supplementary material, which is available to cordingly, plants use a variety of different mecha- authorized users. nisms to minimize the costs of nectar production, Corresponding author: M. Cavalcante, including variation in secretion patterns, sugar [email protected] composition, and accessibility of nectaries Manuscript editor: Klaus Hartfelder (Lovett-Doust and Lovett-Doust 1988). 506 M. C. Cavalcante et al. Nectar secretion patterns, effects of nectar re- main reward offered by this plant species. Nectar moval, and factors determining the chemical con- secretion patterns and floral structures that present stituents of floral nectar, as well as strategies to nectar play important roles in determining plant- protect the floral rewards, have been previously pollinator relationships, particularly in relation to studied in many angiosperm species (e.g., Galetto spatio-temporal visitation patterns and the pollina- and Bernardello 1992; Stiles and Freeman 1993; tion efficacies of different flower visitors. Thus, a Van Wyk 1993; Torres and Galetto 1998; Navarro better understanding of the relationships between 1999; Bernardello et al. 1999; Goulson 1999; pollinators and flowers of Lecythidaceae, especial- Galetto et al. 2000; Galetto and Bernardello ly those only offering nectar as a reward, requires 2003), because all these processes are essential to intensive study of visiting pollinator species understand plant-pollinator interactions and their (Knudsen and Mori 1996), their visitation patterns, implications for plant reproductive success and the study of temporal patterns in nectar pro- (Cruden et al. 1983; Galetto and Bernardello duction (e.g., volume, composition, concentration) 1992). For example, some plant species open their (Galetto and Bernardello 2004, 2005). flowers full of nectar (e.g., Rivera et al. 1996; The Brazil nut (B. excelsa ) is a large-sized Guerra et al. 2014) and others with little nectar arborous plant species which grows in Amazon inside (Galetto and Bernardello 1992, 1993; forests of Brazil, Bolivia, Colombia, Guiana, Su- Amorim et al. 2013), affecting the foraging behav- riname, Peru, and Venezuela at a natural density of ior of their floral visitors (Real 1981; Roubik et al. 1.3 trees/ha (Peres and Baider 1997;Wadtetal. 1995). Besides that, distinct groups of floral visitors 2005). Income generated from the harvest and show preferences according to nectar characteris- natural collection of its seeds (Brazil nuts) is of tics such as sugar concentration or composition. paramount importance for socioeconomic devel- For example, although bee-pollinated plant species opment in these regions (Wadt et al. 2005). usually present hexose-predominant or hexose-rich B. excelsa flowers are hermaphrodite and poten- nectars in their flowers (> 75% of studied plant tial pollinators start to visit its flowers for nectar species), large bees (e.g., Euglossini, Centridini) from the moment of their opening (Prance and prefer visiting flowers that provide sucrose-rich Mori 2004; Santos and Absy 2012). The Brazil nectar with high sugar concentrations (35–50%) nut tree is predominantly xenogamous and the (Roubik et al. 1995; Baker and Baker 1983; main pollinators are known to be large bees Galetto and Bernardello 2003). (Prance 1976;Morietal.1978;Maués2002; Odors seem to be a secondary attractant in Cavalcante et al. 2012). However, among the ten Neotropical species of Lecythidaceae, of which genera of New World Lecythidaceae, flowers of many species are bee-pollinated and produce nec- B. excelsa show the most complex morphologies tarasreward(35–39% sugar concentration and that greatly restrict access to nectar by many 10–60 μL/flower, depending on the species; flower-visiting insects (Prance 1976;Morietal. Knudsen and Mori 1996). Floral scent compounds 1978;MoriandPrance1981; Tsou and Mori in nectar-rewarding Lecythidaceae could indicate 2007; Potascheff et al. 2014). Flowers bear a that male euglossine bees may be one of the main chamber made of congruent staminoids that form pollinators, although this does not necessarily pre- a robust structure (hood or helmet) that covers the clude visitation by female euglossine bees reproductive structures (stamens and stigma) and (Knudsen and Mori 1996). nectaries. B. excelsa produces fruits and seeds by Studies on Lecythidaceae nectar are rare. both xenogamous and geitonogamous pollination, Potascheff et al. (2014) focused on the nectar but under cultivation, the species suffers from volume and concentration of Eschweilera nana , severe pollen limitation (Cavalcante et al. 2012). while Freeman et al. (1991) observed that the Asi- The aims of the present study were to deter- atic species Barringtonia asiatica presents noctur- mine nectar sugar composition and nectar secre- nal flowers which secrete nectar containing 97.6% tion dynamics throughout the anthesis of sucrose.