Spatial Distribution of a Population Of... 1
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Spatial distribution of a population of... 1 SPATIAL DISTRIBUTION OF A POPULATION OF Pentaclethra macroloba (Willd.) KUNTZE IN A FLOODPLAIN FOREST OF THE AMAZON ESTUARY1 Adelson Rocha Dantas2*, Luiz Carlos Marangon3, Marcelino Carneiro Guedes4, Ana Lícia Patriota Feliciano3 and Ana Claudia Lira-Guedes4 1 Received on 25.05.2015 accepted for publication on 21.06.2017. 2 Instituto Nacional de Pesquisas da Amazônia, Programa de Pós-Graduação em Ecologia, Manaus – Brasil. E-mail: <[email protected]>. 3 Universidade Federal Rural de Pernambuco, Departamento de Ciência Florestal, Recife – Brasil. E-mail: <[email protected]> and <[email protected]>. 4 Empresa Brasileira de Pesquisa Agropecuária, Centro de Pesquisa Agroflorestal do Amapá , Macapá – Brasil. E-mail: <[email protected]> and <[email protected]>. *Corresponding author. ABSTRACT – We analyzed the spatial distribution pattern of Pentaclethra macroloba to assist in the development of management plans and conservation of this species in estuarine floodplain forests. The study was conducted in an area of floodplain forest of 55.94 ha, in Mazagão, Amapá, Brazil. A census was performed and all individuals of > 30 cm circumference at breast height (CBH), living and dead, were georeferenced. Ripley’s K function was used to analyze the spatial distribution of the population, including mortality and diameter classes. The relationships between distance to the edge of inundation and density and basal area of the population were analyzed using linear regression. We recorded 993 live individuals and 22 dead. The distribution of the population was found to vary with the topographic gradient associated with the distance to the edge of the inundation. Diameter classes 1 (< 13.2 cm) and 2 (< 18.6 cm) were spatially aggregated close to the water and randomly distributed far from the water. Classes 3 (< 24.0 cm) and 4 (< 29.5 cm) were aggregated at all distances, and classes 5 (< 34.9 cm), 6 (< 40.3 cm), 7 (< 45.7 cm) and 8 (< 51.2 cm), were randomly distributed at all distances. Mortality was randomly distributed. The population density was 18 trees ha-1 with a basal area of 49 m² ha-1, and both density (r2=0.51; p=0.0195) and basal area (r2=0.67; p=0.0039) increased with the distance from the edge of the inundation. We conclude that the spatial distribution and successful establishment of this species in floodplain forest are related to the dynamics of flooding and the resultant impacts on dispersal processes. Keywords: Autoecology of oil bean tree; Ripley’s K function; Dispersal. DISTRIBUIÇÃO ESPACIAL DE UMA POPULAÇÃO DE Pentaclethra macroloba (Willd.) KUNTZE EM FLORESTA DE VÁRZEA DO ESTUÁRIO AMAZÔNICO RESUMO – Analisou-se o padrão de distribuição espacial de Pentaclethra macroloba para auxiliar em planos de manejo e conservação desta espécie nas florestas de várzeas estuarinas. O estudo foi realizado em floresta de várzea de 55,95 ha, na cidade de Mazagão, Amapá. Foi realizado um censo e georreferenciamento de todos os indivíduos, vivos e mortos, que apresentavam a Circunferência na Altura do Peito (CAP) > 30 cm. A Função K de Ripley foi utilizada para analisar o padrão de distribuição populacional, da mortalidade e das classes diamétricas. A relação da distância de inundação com a densidade e a área basal da população foi analisada utilizando regressão linear. Foram inventariados 993 indivíduos vivos e 22 mortos. A população obteve padrão agregado ao longo do gradiente topográfico de inundação. As classes de diâmetro 1 (< 13,2 cm) e 2 (< 18,6 cm) iniciaram com padrão agregado, finalizando com padrão aleatório. As classes 3 (< 24,0 cm) e 4 (< 29,5 cm) obtiveram padrão totalmente agregado e nas classes seguintes, 5 (< 34,9 cm), 6 (< 40,3 cm), 7 (< 45,7 cm) e 8 (< 51,2 cm) o padrão foi aleatório. O padrão da mortalidade foi aleatório. A densidade Revista Árvore. 2017;41(4):e410406 http://dx.doi.org/10.1590/1806-90882017000400006 2 DANTAS AR et al. populacional foi 18 indivíduos ha-1 e área basal de 49 m² ha-1 dos indivíduos vivos. A densidade (r2=0,51; p = 0,0195) e a área basal (r2 = 0,67; p = 0,0039) dos pracaxizeiros aumentam com o afastamento do gradiente de distância do rio. A dinâmica de inundação e o processo de dispersão são fatores relacionados com o padrão espacial e o sucesso no estabelecimento da espécie. Palavras-chave: Autoecologia de pracaxi; Função K de Ripley; Dispersão. 1. INTRODUCTION spikes, with 200 bisexual flowers per stem. The fruits are dry, dehiscent green pods that turn brown when High rainfall and climatic seasonality lead to periodic ripe. The seeds are dark brown and opaque with flooding of areas along the main river systems of the superficial depressions forming prominent lines that Amazon (Wittmann et al., 2006), with areas flooded are sometimes reticulated close to the base of the by white water being known as várzeas or floodplain seed (Flores, 2003). Flowering occurs between July forests (Wittmann et al., 2010). Floodplain forests are and September, the dry season, and fruits mature energetically open systems that contain large volumes and seeds are dispersed between January and April, of freshwater and sediments, and are driven by daily the period of highest rainfall (Freitas et al., 2003). (Rabelo, 2000) and annual (Junk, 1989) cycles of flooding. The primary dispersal mechanism is autochory, with The flood pulse is unimodal and predictable, leading seeds dispersed by explosive dehiscence (Flores, to a terrestrial and an aquatic phase (Junk, 1997). Due 2003). Beyond this, in the Amazonian floodplains to constant flooding, a large amount of sediment, P. macroloba has hydrochory as a secondary dispersal originating in the Andes or pre-Andes (Mcclain and mechanism (Williamson and Costa, 2000). Naiman, 2008), is carried to these areas (Rabelo, 2000) Approximately 90% of seeds germinate (Flores, 2003). which leads to highly fertile soils (Rabelo, 2000). The species is a pioneer (Condé and Tonini, 2013), Floodplain forests play an important role in the organic occurring in high densities in the Amazon Estuary carbon balance in the Amazon Basin, are areas of high (Carim et al., 2017), and being characterised by rapid rates of productivity of aquatic plants and are important growth and a high capacity for regeneration (Sears sources of methane for the troposphere (Melack et and Pinedo-Vasquez, 2005). The bark of this species al., 2009). is used to dye the skin and also as a medicine to These forests contain an important diversity of treat dysentery and diarrhoea (Cordero and Boshier, trees, with approximately 1,000 tree species tolerant 2003), and the leaves can be used to treat fungal to the patterns of flooding already recorded from the infections (Shebitz et al., 2013). However, its greatest Amazon Basin (Wittmann et al. 2006). These trees are potential is in the oil extracted from its seeds, that an important source of resources for a majority of the has medicinal properties (Pesce, 2009; Costa et al., rural population of the Amazon (Wittmann et al., 2010), 2013) that could make it very attractive to the cosmetics providing a variety of timber and, most importantly, and herbal medicine industries, although it is not non-timber forest products (NTFPs) that are sources yet marketed on a large-scale. of food, pharmaceutics and cosmetics. These NTFPs Ecological studies that aim to investigate the include: fruits (taperebá, urucuri), oils (andiroba, pracaxi), spatial distribution of this species within the forest barks (andiroba, ucuúba, pracaxi), resins (breu-branco), are fundamental for programmes of management and and latex (seringueira), etc. conservation. According to Condit et al. (2000), the One species of particular importance as a source spatial distribution of a species allows for an of NTFPs is Pentaclethra macroloba (Willd.) Kuntze. understanding of both reproductive processes and This species belongs to the family Fabaceae and is how the resources available in the community are popularly known in the Amazon region as “pracaxi”. used. Indeed, understanding how individuals of P.macroloba reaches a height of approximately 14 m a species distribute themselves in a community is and a DBH (Diameter at Breast Height) of up to 59 one of the first steps to understanding forest dynamics cm (Projeto Florestam). The leaves are bipinnate. The (Hay et al., 2000), natural history and ecological inflorescences are in the form of terminal or subterminal interactions between populations (Forget et al., 1999). Revista Árvore. 2017;41(4):e410406 Spatial distribution of a population of... 3 The objective of this study was to analyse the The soils of the area are classified as Melanic spatial distribution pattern of P. macroloba to aid in Gleysols (Embrapa, 2013), soils that have a medium the development of management plans and in the species’ texture, poor drainage and medium to high natural fertility conservation, aiming to support sustainable exploitation (Yokomizo et al., 2010). According to surveys by of this species in estuarine floodplain forests. pedologist Raimundo Cosme Oliveira Júnior (Embrapa Amazônia Oriental) carried out in the project “ecologia 2. MATERIAL AND METHODS e manejo para uso múltiplo de florestas de várzea do estuário amazônico - Florestam”, all the soils of this 2.1. Characterization of the study area region of the estuarine floodplain are classified as Melanic The study was conducted in the “Campo Experimental Gleysols Ta Eutrophic, with a predominance of silt do Mazagão - CEM”, owned by Embrapa Amapá. The CEM textures. Both the sediment and the soil are formed is located in the municipality of Mazagão, in the south of by smectite, ilite, kaolinite, goethite, anatase and quartz the state of Amapá, at 00°02’33" south and 51°15’24" west, (unpublished data). and at an altitude of 15 m (Yokomizo et al., 2010). It has 2.2.