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Sociobiology 64(2): 191-201 (June, 2017) DOI: 10.13102/Sociobiology.V64i2.1232

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Sociobiology 64(2): 191-201 (June, 2017) DOI: 10.13102/Sociobiology.V64i2.1232 View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by Portal de Periódicos Eletrônicos da Universidade Estadual de Feira de Santana (UEFS) Sociobiology 64(2): 191-201 (June, 2017) DOI: 10.13102/sociobiology.v64i2.1232 Sociobiology An international journal on social insects RESEARCH ARTICLE - BEES Male Orchid Bees (Hymenoptera: Apidae: Euglossini) in Canopy and Understory of Amazon Várzea Floodplain Forest. I. Microclimatic, Seasonal and Faunal Aspects PS Vilhena¹, LJ Rocha², CA Garófalo¹ 1 - Faculdade de Filosofia, Ciências e Letras de Ribeirão Preto, Universidade de São Paulo (USP), Ribeirão Preto-SP, Brazil 2 - Universidade Federal do Amapá, Macapá-AP, Brazil Article History Abstract Floodplain forests are important ecosystems of the Amazon Basin. The aim Edited by of the present study was to investigate whether male orchid bees have Og de Souza, UFV, Brazil Received 01 November 2016 preference for arboreal strata in a forest with a dynamic understory. Traps Initial acceptance 07 March 2017 with aromatic scents to attract bees were placed at heights of 25 m and 1.5 Final acceptance 01 May 2017 m. Data loggers were used to register temperature and relative humidity Publication date 21 September 2017 at both heights. A total of 835 individuals belonging to 28 species were sampled, 645 males of 23 species in the understory and 190 males of 24 Keywords 0 Amapá, vertical stratification, diversity, species in the canopy. The temperature was 1.4 C higher in the canopy and euglossine bees. the air relative humidity was 11.58% higher in the understory. Temperature, relative humidity and winds were the most important abiotic variables Corresponding author influencing the assemblage in the canopy. The abundance increased in the Patricia dos Santos Vilhena understory and decreased in the canopy in the less rainy season. Euglossa Laboratório de Ecologia e Evolução de cognata Moure, Eulaema meriana Olivier and Euglossa ignita Smith were Abelhas e Vespas Av. Bandeirantes, 3900, Monte Alegre species closely associated with strata and seasons. The present study revealed CEP 14040-901, Ribeirão Preto higher richness and abundance in the understory all months of the year, São Paulo, Brasil. reinforcing the likelihood that orchid bees forage preferably in the lower E-Mail: [email protected] layer of rainforests. Introduction found that tree species pollinated by small bees were especially common in the understory, whereas species pollinated by Studies have shown significant differences in the medium-sized to large bees were predominant in the canopy of a diversity and community composition of insects across distinct tropical forest. Roubik (1993) performed systematic collections arboreal strata, such as canopy and understory, regardless of of bees from the canopy and understory in two forests in Panama the taxon and type of forest analyzed (Smith, 1973; Dulmen, and found that two species of nocturnal bees were conspicuously 2001; Giovanni et al., 2015). These differences may be abundant in the canopy, despite of that most of the species did related to abiotic conditions and biotic resources, such as light not have a preference for a particular stratum. Sobek et al. (2009) availability, temperature, wind exposure, forest structure and reported spatial stratification in the abundance of cavity-nesting composition, food and nesting sites, which differ in intensity bees and wasps and their parasitoids as well as higher parasitism and availability between strata (Ulyshen, 2011). For some rates in the canopy than the understory in temperate deciduous groups of insects these differences are fairly important so that forests in Germany. In a study on the vertical distribution of trap- there are species restricted to specific strata (Erwin, 1983). nesting bees along a fragmentation gradient of secondary forest As emphasized by Ulyshen et al. (2010), few studies remnants in Costa Rica, Stangler et al. (2016) found that bees were have been conducted to determine the occurrence of the vertical more abundant in the canopy and understory in comparison to an distribution of bees. Among such studies, Bawa et al. (1985) intermediate height but the diversity was higher in the canopy. Open access journal: http://periodicos.uefs.br/ojs/index.php/sociobiology ISSN: 0361-6525 192 PS Vilhena, LJ Rocha, CA Garófalo – Male orchid bees in strata of várzea floodplain forest Some authors have investigated the occurrence of brasiliensis Muell. Arg., Euphorbiaceae), andiroba nut (Carapa vertical stratification in populations of orchid bees. Roubik guianensis Aubl., Meliaceae), pracuúba (Mora paraensis (1993) found that large euglossine bees demonstrated a tendency Ducke, Leguminosae-Caesalpinioideae), palm species, such as to forage high, which was directly related to their capacity for heat açaí (Euterpe oleracea Mart., Arecaceae) and others (Prance, loss during flight, compared to smaller euglossine bees. Oliveira 1979; Carim et al., 2017). This vegetation gives to the forest and Campos (1996), in two areas of Terra Firme forest in the an over-20-m canopy in height and a clear understory that is Amazon, found similar values of richness and abundance flooded up to 2 m by tides mainly from the Amazon River and between canopy and understory, but higher diversity in the some creeks (igarapés), such as Paxicú and Fortaleza, that flow canopy; the authors also reported that some species were more throughout the forest (Drummond et al., 2008). associated with the canopy and others were more common The climate is typical of Tropical Forest, Af according in the understory. In areas of the Atlantic Rainforest (coastal to Köppen’s classification (Peel at al., 2007), with high rainfall Brazil), Martins and Souza (2005) found that the abundance throughout most of the year. Mean annual precipitation is and richness of euglossine bees were higher in absolute approximately 2100 mm. A rainier season extends from January numbers in the understory, but the diversity was higher in the to July and a less rainy season extends from August to canopy. Studying population fluctuations of euglossine bees December. Mean annual temperature is about 27 ºC and mean in a remnant of Cerrado scrubland in the state of Mato Grosso annual humidity is higher than 78% (Drummond et al., 2008). do Sul, Brazil, Ferreira et al. (2011) found no differences in species richness or diversity between strata, but the abundance Sampling of males was higher in the canopy than the understory. All these studies carried out with orchid bees had focused on The fieldwork was carried out once a month, from differences and similarities in the richness, abundance and January 2011 to March 2012, in an open natural area 500 m from diversity between canopy and understory, but none of them the edge of the forest. Twenty-six traps were used to capture the assessed the influence of abiotic conditions on the community bees. Each trap consisted of a 2-L plastic bottle with four holes and distribution of species across the different strata. measuring 3 cm in diameter spaced 5 cm apart and 17 cm up Therefore, the aim of the present study was to investigate from the bottom of the bottle. Transparent hoses measuring 4 cm how the assemblage of male orchid bees is structured in canopy in length were cut in half longitudinally and attached to the holes and understory of a várzea floodplain forest in the Brazilian to facilitate the entrance of the males. Three cotton swabs with Amazon, taking into consideration that floodplains have a more extra cotton (ball diameter: 2 cm) were contained in each trap and dynamic understory than non-flooded forests, which could favor impregnated with an aromatic scent. Thirteen scents were used: faunal differences in both strata. Microclimatic differences 1.8-cineole, eugenol, vanillin, benzyl acetate, methyl salicylate, regarding temperature and air relative humidity were analyzed methyl cinnamate, β-ionone, benzyl benzoate, β-myrcene, ethyl in each stratum and the influence of such abiotic variables, butyrate, linalool, methyl benzoate and 2-phenyletanol. One pair together with others as rainfall, wind speed and tidal height, was of traps with the same scent was securely fixed, to avoid wind investigated on the assemblage. shaking, on the same tree at the height of 25 m (canopy) and 1.5 m above the ground (understory). The trees were approximately 10 Material and Methods m one from each other, encompassing an area of approximately 150 m². The traps were monitored from 8:00 a.m. to 6:00 p.m., Study area being replenished every four hours, when the attracted males were removed. This study was conducted in the Protected Area of Fazendinha, category V of the IUCN list (Dudley, 2008), which Abiotic variables is located in the southeastern portion of the state of Amapá, Eastern Amazon, Brazil (00º03’02”S 051º07’51” W). The area General abiotic data of the area: rainfall (mm), temperature encompasses 137 ha comprised mostly of dense tidal várzea (ºC), air relative humidity (%) and wind speed (m/s) were forest (Prance, 1979), a kind of floodplain subject to floods by obtained from the Núcleo de Hidrometereologia e Energias white-water rivers, which is different of igapó, another kind of Renováveis/ Instituto de Pesquisas Científicas e Tecnológicas Amazonian floodplain subject to floods by black or clean-water do Estado do Amapá (NHMET/ IEPA). Microclimatic data of rivers (Prance, 1979). These forests represent about 27% of the each arboreal stratum [temperature (ºC) and air relative humidity Amazon Basin and differ one from each other regarding many (%)] were obtained from two climatic data loggers (HOBO® aspects, as forest structure and composition, number of arboreal Pro Series) placed at 25 m and 1.5 m in height to record data strata, height and architecture of the canopies, density coverage, on an hourly basis. These loggers were used from April 2011 soil composition, and so on (Pires & Prance, 1985; Piedade et to March 2012 only on sampling days.
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