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Germination and Allometry of the Native Palm Tree Euterpe Edulis Compared to the Introduced E

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Germination and Allometry of the Native Palm Tree Euterpe Edulis Compared to the Introduced E Germination and allometry of the native palm tree Euterpe edulis compared to the introduced E. oleracea and their hybrids in Atlantic rainforest Tiberio, FCS.a*, Sampaio-e-Silva, TA.a, Dodonov, P.a, Garcia, VA.b and Silva Matos, DM.a aLaboratório de Ecologia e Conservação, Departamento de Botânica, Universidade Federal de São Carlos – UFSCar, Rod. Washington Luís, Km 235, SP 310, São Carlos, SP, Brazil bPolo Regional de Desenvolvimento Sustentável dos Agronegócios do Vale do Ribeira, Agência Paulista de Tecnologia dos Agronegócios, SAA-SP, Rod. Regis Bittencourt, Km 460, Pariquera-Açu, SP, Brazil *e-mail: [email protected] Received November 30, 2011 – Accepted February 14, 2012 – Distributed November 30, 2012 (With 1 figure) Abstract Palms are distinctive plants of tropics and have peculiar allometric relations. Understanding such relations is useful in the case of introduced species because their ability to establish and invade must be clarified in terms of their responses in the new site. Our purpose was to assess the survival and invasive capacity of an introduced palm species in the Atlantic rainforest, Euterpe oleracea Mart., compared to the native Euterpe edulis Mart. and to the hybrids produced between the two species. Considering this, we compared the allometry in different ontogenetic stages, the germination rates, and aspects of the initial development. The ontogenetic stages proposed for both Euterpe illustrated the growth patterns described for palm trees. E. oleracea and hybrids adjusted to the geometric similarity allometric model, while E. edulis presented a slope greater than would be expected considering this model, indicating a greater height for a given diameter. E. oleracea showed the same amount of pulp per fruit as E. edulis and a similar initial development of seedlings. The main differences observed were a lower germination rate and a faster height gain of E. oleracea seedlings. We conclude that E. oleracea, which is similar to E. edulis in aspects of allometry, development, seed and seedling morphology, may be an important competitor of this native palm tree in the Atlantic Forest. Keywords: palms, hybridization, growth model, species introduction. Germinação e alometria de Euterpe edulis comparadas à palmeira introduzida Euterpe oleracea e seus híbridos na floresta Atlântica Resumo Palmeiras são plantas características dos trópicos que apresentam relações alométricas peculiares. Compreender tais padrões pode ser útil no caso de espécies introduzidas, uma vez que sua habilidade de estabelecimento e invasão deve ser esclarecida em relação as suas respostas à nova localidade. Nosso propósito foi compreender a sobrevivência e a capacidade de invasão de uma palmeira introduzida na floresta Atlântica, Euterpe oleracea Mart. (açaizeiro) comparada à palmeira nativa Euterpe edulis Mart. (juçara), também considerando seus híbridos. Para isso comparamos suas relações alométricas em estádios ontogenéticos, sua germinação e seu desenvolvimento inicial. Os estádios ontogenéticos propostos para ambas as Euterpe ilustraram os padrões de crescimento esperados para palmeiras. E. oleracea e híbridos apresentaram-se sob o modelo alométrico de similaridade geométrica e E. edulis apresentou inclinação maior do que este modelo. E. oleracea produziu a mesma quantidade de polpa por fruto que E. edulis. As principais diferenças observadas foram menor taxa de germinação e maior velocidade de crescimento em altura para as plântulas de E. oleracea. Em conclusão, nossos resultados indicam que E. oleracea, sendo similar a E. edulis em termos de alometria, desenvolvimento e morfologia de sementes e plântulas, pode ser um importante competidor para a espécies nativa na Floresta Atlântica. Palavras-chave: palmeiras, hibridização, modelo de crescimento, introdução de espécies. Braz. J. Biol., 2012, vol. 72, no. 4, p. 955-962 955 Tiberio, FCS. et al. 1. Introduction Considering this, we compared their ontogenetic development, allometry, seed and seedling characteristics and germination. Palms are typical exponents of tropical plants and Since the two species have similar morphology when play an important role in natural and human communities occurring in their natural ranges (Henderson et al., 1995), (Henderson et al., 1995; Tomlinson, 2006). They have we expected similar allometry and germination rates when impressive development and and great ecological importance, co-occuring in the same environment. This would indicate especially as food resources. After their fruit are eaten and the potential competition between the two species and their seeds land on the ground, development must start the potential of E. oleracea to invade the Atlantic forest. quickly because most of them do not form soil seed banks (Henderson et al., 1995). Palms are also unique in the way 2. Methods they build long, tall lived trees entirely by primary growth, i.e., all tissues result directly from the activity of shoot Study area: We sampled the populations of Euterpe edulis, and apical meristems, ensuring minimum susceptibility to E. oleracea and their hybrids in Carlos Botelho State Park fire, pathogens and wind damage (Tomlinson, 2006). This (CBSP), located in south-eastern Brazil, with a total area growth results in a unique allometric pattern, where height of 37,644 ha and the altitudes between 20 and 1000 m may increase almost unlimitedly by the plant diameter (SÃO PAULO, 2008). The vegetation is a typical tropical when compared to other tree species (Rich, 1987). rainforest with Af climate (sensu Köppen 1948) and Allometric patterns of plants result from the balance average rainfall from 1700 to 2400 mm (SÃO PAULO, of investments in support structures and photosynthetic 2008). In 1970, E. oleracea was planted in a floodplain biomass (Niklas, 1995; Niklas, 2004). Aerial parts of area within the Park. a plant are submitted to forces like gravity and wind, Seeds were collected from plants located in the so they have to allocate energy to support themselves experimental plantation of Agência Paulista de Tecnologia while growing, and must also provide photosynthetic dos Agronegócios (APTA) (Agrarian Technology Agency structures to ensure biomass assimilation. Allometric of São Paulo State) in the municipality of Pariquera- relationships may change during the development of Açu, São Paulo, south-eastern Brazil. The objective of plant individuals, with each ontogenetic stage presenting these E. oleracea plantations was to produce hybrids a different allometry between height and diameter, as with E. edulis to improve the development of palm heart a response to dissimilar environmental, morphological (Bovi et al., 1987a). and physiological conditions experienced by individuals Study species: the jussara palm, E. edulis, occurs naturally from different stages (Gatsuk et al., 1980; Weiner, 2004). in the Atlantic rainforest on the Brazilian coast and may It is already recognized that non-indigenous species are also be found in Paraguay and Argentina (Henderson et al., intensively dispersed for economic purposes (Levine et al., 1995). It has an arborescent habit, with a single stem, and 2003), and this applies to many palm trees (Holmquist et al., may reach 15 m in height in the study site. It produces 2010; Henderson et al., 1995; Svenning, 2002; Meyer et al., globular purple-black fruit with 1-1.4 cm in diameter 2008). Although the establishment of plant populations (Henderson et al., 1995). This species has been under outside their natural range of occurrence is not common threat due to indiscriminate exploitation, mostly of which (Richardson et al., 2000), some of these populations can become successfully established (Mooney and Cleland, is illegal, since the second half of the twentieth century 2001) and modify ecosystem functions, as well as the (Silva Matos and Bovi, 2002). As it produces a single availability or quality of resources and space (Levine et al., stem and does not resprout, the plant dies when the palm 2003). Therefore, assessment of morphological traits can heart is extracted. Currently, the presence of E. edulis is also be a tool for evaluating the status of non-indigenous restricted to remote areas of the Atlantic rainforest (Silva species, since phenotypic plasticity indicates its ability Matos and Bovi, 2002). to adapt to different environments (Tucic et al., 2006). The assaí palm, E. oleracea, is commonly found Amongst many examples from Brazil, two ecological along the Amazon river basin in the North of Brazil, and and economically important palm trees with dissociated therefore it is geographically distant from E. edulis in the occurrence are now being cultivated in the same Atlantic rainforest (Henderson et al., 1995). E. oleracea environments: Euterpe oleracea, originating from the produces multiple stems and can reach 12-20 meters in Amazon rainforest, has been cultivated on a large scale in height. Its apical meristem is also exploited to produce the Atlantic Rainforest, the natural habitat of the congener palm heart (Henderson et al., 1995). It bears purple-black palm E. edulis, since the late 1970s. Fruit from both species globular fruits with 1-2 cm in diameter (Henderson et al., is important in terms of food resources for birds in their 1995; Lorenzi et al., 2004) which are harvested for assaí original areas (Galetti et al., 1999; Moegenburg and Levey, cream production. The palm heart from E. oleracea has 2003; Genini et al., 2009). When co-existing, these palms a lower quality than its congener but, as a plant with produce hybrids (Bovi et al., 1987a; Campos et al., 1991). multiple stems, the production may be higher and the plant The objective of this study was to assess the phenotypic is not necessarily killed during palm heart extraction as plasticity of the introduced species E. oleracea and its occurs with E. edulis. The better quality of its palm heart, hybrids with the native E. edulis in the Atlantic rainforest. the increasing demand of the consumer market and the 956 Braz. J. Biol., 2012, vol. 72, no. 4, p. 955-962 Germination and allometry of E. edulis, E. oleracea and hybrids scarcity of E. edulis made it one of the most expensive a is the slope value that represents the proportionality palm hearts in Brazil.
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