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Floral Biology, Morphology and Ecological Niche Modelling Of Bol. Mus. Para. Emílio Goeldi. Cienc. Nat., Belém, v. 9, n. 3, p. 465-479, set.-dez. 2014 Floral biology, morphology and ecological niche modelling of Caraipa grandifolia (Calophyllaceae), an important Amazonian floodplain tree Biologia floral, morfologia e modelagem ecológica de nicho de Caraipa grandifolia (Calophyllaceae), uma árvore importante das planícies inundáveis amazônicas Leonardo de Melo VersieuxI, André Luis AcostaII, Alexandre Luis JordãoIII, Alexsandro ZidkoIV, Ulysses Madureira MaiaV IUniversidade Federal do Rio Grande do Norte. Natal, Rio Grande do Norte, Brasil IIUniversidade de São Paulo. São Paulo, São Paulo, Brasil IIIInstituto de Pesquisas Científicas e Tecnológicas do Estado do Amapá. Macapá, Amapá, Brasil IVInstituto Nacional de Pesquisas da Amazônia. Manaus, Amazonas, Brasil VUniversidade Federal Rural do Semi-Árido. Mossoró, Rio Grande do Norte, Brasil Abstract: The present work aims to investigate the floral biology and morphology of one member of the Calophyllaceae (or Clusiaceae sensu lato), Caraipa grandifolia in the Caxiuanã National Forest, in the eastern Brazilian Amazon and aims to identify the bioclimatic affinities of this species, providing insights about its environmental requirements and mapping its potential distribution, given the paucity of data about flood-tolerant trees. Nine flowering individuals were located along a 2 km stretch of the Anapu river, and were visited by eleven different species (some of which are potential pollinators) including cockroaches, bees, ants, and wasps. Flower anthesis occurred between 05:00 and 07:30. The flowers are odourous, and osmophores were observed along the petals and also on top of each individual anther. The main reward collected by bees was pollen and an oily resin/scent produced by anther glands, while four different ant species collected nectar in inconspicuous extranuptial nectaries along the flower pedicel, a morphological feature previously unreported for the genus. Caraipa grandifolia is considered to be an important tree species in the flooded forest ecosystems given the number of interactions with which it is involved and its potentially wide distribution indicated by our modelling. Keywords: Clusiaceae. Dictyoptera. Igapó. Resumo: O presente trabalho tem como objetivo investigar a biologia floral e morfologia de uma espécie de Calophyllaceae (ou Clusiaceae sensu lato), Caraipa grandifolia, na Floresta Nacional de Caxiuanã, na Amazônia oriental brasileira, e identificar as afinidades bioclimáticas dessa espécie. Fornecemos dados sobre suas exigências ambientais e mapeamos a sua distribuição potencial, dada a escassez de informações sobre árvores tolerantes ao alagamento. Nove indivíduos floridos foram localizados ao longo de um trecho de 2 km do rio Anapu e foram visitados por onze espécies diferentes (algumas são potenciais polinizadores), incluindo baratas, abelhas, formigas e vespas. A antese floral ocorreu entre 05:00 e 07:30 h. As flores são odoríferas, e osmóforos foram observados ao longo das pétalas e também no topo de cada antera. As principais recompensas obtidas pelas abelhas são pólen e resina oleosa com perfume produzida por glândulas nas anteras, enquanto quatro espécies diferentes de formigas coletaram néctar em nectários extranupciais discretos, até então desconhecidos para o gênero, localizados ao longo do pedicelo da flor. Caraipa grandifolia é considerada uma importante espécie arbórea nos ecossistemas florestais inundados, dado o número de interações em que está envolvida e a sua distribuição em potencial, indicada na modelagem. Palavras-chave: Clusiaceae. Dictyoptera. Igapó. VERSIEUX, L. M., A. L. ACOSTA, A. L. JORDÃO, A. ZIDKO & U. M. MAIA, 2014. Floral biology, morphology and ecological niche modelling of Caraipa grandifolia (Calophyllaceae), an important Amazonian floodplain tree. Boletim do Museu Paraense Emílio Goeldi. Ciências Naturais 9(3): 465-479. Autor para correspondência: Leonardo de Melo Versieux. Universidade Federal do Rio Grande do Norte. Centro de Biociências. Departamento de Botânica e Zoologia. Campus Universitário. Natal, RN, Brasil. CEP 59078-900 ([email protected]). Recebido em 27/11/2013 Aprovado em 14/11/2014 Responsabilidade editorial: Alexander C. Lees 465 Floral biology, morphology and ecological niche modelling of Caraipa grandifolia (Calophyllaceae), an important Amazonian floodplain tree INTRODUCTION The genus Caraipa includes 30 species in South Despite the important role played by tropical forests in America of which 22 occur in Brazil, varying from large maintaining global climate and carbon cycles (e.g. Bonan, trees to shrubs and occurring in many distinct vegetation 2008), both the biodiversity and underlying ecological types (Gustafsson & Nee, 2004; Stevens, 2007; Bittrich et processes in the Brazilian Amazonian forest are still poorly al., s. d.). This genus is traditionally positioned within the understood (e.g. Feeley & Silman, 2009; Hopkins, 2007; Clusiaceae or, more recently, within the Calophyllaceae Hubbell et al., 2008; Milliken et al., 2010). Knowledge of (Ruhfel et al., 2011; Souza & Lorenzi, 2012; Stevens, 2001 the biology of the flood-tolerant trees of the Amazonian - onwards). Members of the Calophyllaceae are often floodplain is particularly poor (Parolin, 2012). The considered important timber species (e.g. Calophyllum eastern Brazilian Amazon lies astride the Amazonian brasiliense, Guanandi) and are currently represented in Arc of Deforestation (e.g. Vieira et al., 2008) a region Brazil by eight genera and nearly 80 species (Souza & characterised by intense forest degradation from logging Lorenzi, 2012). All species of Caraipa are Neotropical and fires (Nepstad et al., 1999) and concomitant land- and are common forest trees on acidic soils; they have use change due to agriculture (see Adeney et al., 2009). sparse latex, alternate leaves, stellate trichomes and white, Understanding biotic interactions in the remnants of this contorted petals (Kubitzki, 1978; Gustafsson, 2009). rapidly disappearing ecosystem is of utmost conservation Caraipa grandifolia Mart. is a common tree species importance (Gentry, 1988). along the margins of Anapu river inside the Caxiuanã Species distribution modelling (SDM) has been National Forest, Pará State, Brazil. No information was used to investigate the environmental suitability for found in the literature regarding the floral biology and the distribution of discrete species (Hutchinson, 1957; floral visitors of this species, which is popularly known in Soberón, 2010; Soberón & Peterson, 2005). Such the Amazon region as tamaquaré and used as an important models may be useful for species which are distributed medicine ‘óleo de tamaquaré’ to treat different diseases in remote regions of the globe and for which logistical and as a source of wood (Kubitzki, 1978). During our and cost considerations prohibit fieldwork (Brito et al., preliminary field observations we noticed that Caraipa 2009). The Amazon rainforest has several tree species grandifolia inflorescences were constantly visited by various for which basic information about life cycles, biology or species, particularly bees and ants, which motivated us to taxonomy is still incompletely known (e.g. Kubitzki, 1978; proceed on a detailed observation of the floral biology and Parolin, 2012). SDMs have been considered a successful morphology of this Amazonian tree as well as to identify tool to predict the distribution of species in cases like this environmental factors related to occurrence and other (Brito et al., 2009). These techniques combine species suitable areas for C. grandifolia. occurrence data with environmental variables based on a correlative approach to build a representation of the MATERIAL AND METHODS ecological requirements of a species (Anderson et al., This work was carried out in the area surrounding the 2003). Several algorithms have been applied to create Ferreira Penna Scientific Station, in the Caxiuanã National such models, and their computational results can be Forest (hereafter CNF) in Melgaço, Pará State, Brazil projected onto a map showing areas that are similar (1º 42’ 30” S, 51º 31’ 45” W; Figure 1). The selection of to those where the species is known to occur and that flowering adult individuals and observations were carried represents the potential areas of occurrence (Stockwell out using boats. Nine flowering individuals were marked & Peters, 1999; Phillips et al., 2006). in the ‘igapó’ vegetation – flooded portions of the forest 466 Bol. Mus. Para. Emílio Goeldi. Cienc. Nat., Belém, v. 9, n. 3, p. 465-479, set.-dez. 2014 Figure 1. Map of Brazil showing Pará State (A) and the Caxiuanã National Forest (B). Nine blooming individuals of Caraipa grandifolia were mapped along the North and South sides of the Anapu river (C). 467 Floral biology, morphology and ecological niche modelling of Caraipa grandifolia (Calophyllaceae), an important Amazonian floodplain tree along the Anapu river (Figure 1C; points 1 to 9). Individuals (Dafni et al., 2005) we also performed an anatomical study. were monitored in the field from September 17-22 2011. Petals were conserved in 70% ethanol and were later used Temperature and humidity were recorded at the time to obtain transversal sections using a razor blade. This tissue of the observations. Vouchers were deposited in the was bleached by sodium hypochlorite, stained for two Herbarium of the Museu Paraense Emílio Goeldi (MPEG) minutes in Safrablau or Sudan Black 0.02%, mounted as and the Universidade Federal
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