State of Knowledge of Lecythidaceae in Espírito Santo State, Brazil
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BOL. MUS. BIOL. MELLO LEITÃO (N. SÉR.) 36:63-84. OUTUBRO DE 2014 63 State of knowledge of Lecythidaceae in Espírito Santo State, Brazil Michel Ribeiro1,*, Scott Alan Mori2, Anderson Alves-Araújo3 & Ariane Luna Peixoto4 ABSTRACT: Members of Lecythidaceae (Brazil nut family) are usually canopy or emergent trees that dominate in non-flooded forests. The aim of this study was to identify and analyze herbarium specimens and to gather information about the species of Lecythidaceae occurring in the state of Espírito Santo. Collection of specimens of Lecythidaceae in the state began in 1930, with a significant increase in collection effort in the last 20 years. Today, 11 species and two morphospecies have been recorded in Espírito Santo, corresponding to 55% of the species occurring in the entire Atlantic Forest phytogeographic domain, and positioning it as the second state, after Bahia, with the greatest richness in the family in this domain. In Espírito Santo, the group reaches its highest richness in tabuleiro forest, followed by hillside forest and restinga formations. Floristic and phytosociological surveys indicate that the richness and diversity of Lecythidaceae are highest in forests at altitudes between 30 and 200 m a.s.l., and almost all its species occur in low densities. Cariniana ianeirensis is a new record for Espírito Santo. Cariniana ianeirensis, Cariniana legalis, Cariniana parvifolia and Couratari asterotricha are listed as endangered in the Red Book of the Brazilian Flora, and the last two species and Couratari asterophora are among the rarest members of the Brazilian flora. The data demonstrate the importance of Atlantic Forest remnants in Espírito Santo for the conservation of this family of imposing trees. 1 Instituto de Pesquisas Jardim Botânico do Rio de Janeiro. Escola Nacional de Botânica Tropical, Programa de Pós-graduação em Botânica, Rua Pacheco Leão, 2040, Horto, 22460-030, Rio de Janeiro, RJ, Brasil. 2 The New York Botanical Garden, Institute of Systematic Botany, Bronx, 10458-5126, New York, United States. 3 Universidade Federal do Espírito Santo, Centro Universitário Norte do Espírito Santo. Rodovia BR 101 Norte, Km 60 Bairro Litorâneo, 29932-540, São Mateus, ES, Brasil. 4 Instituto de Pesquisas Jardim Botânico do Rio de Janeiro. Rua Pacheco Leão, 915, Horto, 22460-038, Rio de Janeiro, RJ, Brasil. *Author for correspondence: [email protected] Recebido: 7 jul 2014 – Aceito: 9 set 2014 64 RIBEIRO ET AL.: LECYTHIDACEAE IN ESPÍRITO SANTO, BRAZIL Key words: Brazilian Flora, Atlantic Forest, plant conservation, species richness, taxonomy. RESUMO: (Estado do conhecimento de Lecythidaceae no Espírito Santo, Brasil) Espécies de Lecythidaceae (família da castanha-do-Pará) são geralmente árvores de dossel ou emergentes dominantes em florestas não inundáveis. O objetivo deste estudo foi identificar e analisar espécimes de herbário e reunir informações sobre as Lecythidaceae ocorrentes no estado do Espírito Santo. As coleções de espécimes de Lecythidaceae no estado iniciaram em 1930, com aumento significativo do esforço de coleta nos últimos 20 anos. Atualmente, 11 espécies e duas morfoespécies são registradas no Espírito Santo, que corresponde a 55% das espécies ocorrentes no domínio fitogeográfico da Mata Atlântica e posiciona-o como o segundo estado, depois da Bahia, com maior riqueza na família para esse domínio. No estado, o grupo apresenta maior riqueza na floresta de tabuleiro, seguida da floresta de encosta e restinga. Pesquisas florísticas e fitossociológicas apontam maior riqueza e diversidade de Lecythidaceae em florestas com altitudes entre 30 a 200 m e quase todas as suas espécies possuem baixas densidades. Cariniana ianeirensis é uma nova ocorrência para o Espírito Santo. Cariniana ianeirensis, Cariniana legalis, Cariniana parvifolia e Couratari asterotricha são consideradas ameaçadas de extinção pelo Livro Vermelho da Flora do Brasil, as duas últimas espécies mais Couratari asterophora são raras para a flora brasileira. Os dados demostram a importância dos remanescentes de Mata Atlântica no Espírito Santo para a conservação dessa família de arvores imponente. Palavras-chave: Flora do Brasil, Mata Atlântica, conservação de plantas, riqueza de espécies, taxonomia. Introduction The state of Espírito Santo lies entirely within the Atlantic Forest phytogeographic domain. The variations in relief, altitude, temperature, rainfall and soil types across the state contributed to the development of a considerable diversity of ecosystems (Lani et al., 2008). Since the beginning of colonization of the state, the Atlantic Forest has suffered loss of its natural resources due to deforestation for economic reasons. Until the early twentieth century, the original forest covered 65% of the state’s territory; its devastation intensified in the 1960s because of timber exploitation, and especially in the northern region, the state’s forests rapidly declined (IPEMA, 2005; Thomaz, 2010). BOL. MUS. BIOL. MELLO LEITÃO (N. SÉR.) 36. 2014 65 Currently the state has 11% of its original vegetation remaining in scattered and fragmented habitats (Fundação SOS Mata Atlântica & INPE, 2011). Most of the fragments are within privately owned properties. Protected areas include 76 of these fragments, six of which cover more than 10,000 ha (SEAMA, 2014; IPEMA, 2005). Three major vegetation formations occur in Espírito Santo, in three distinct geomorphological provinces (Martin et al., 1993): (1) hillside forest, located in mountainous areas on Precambrian crystalline rocks (Assis, 2007; Assis et al., 2007; Saiter et al., 2011); (2) tabuleiro forest (coastal lowland forests), on flat to gently sloping landscapes of the Tertiary tablelands composed of sediments of the Barreiras Formation (Jesus & Rolim, 2005; Simonelli, 2007; Peixoto et al., 2008), and (3) restinga vegetation (coastal dune scrub forest), developed on the sandy sediments of the Quaternary coastal plain (Fabris & Cesar, 1996; Pereira, 2007; Giaretta et al., 2013). Members of Lecythidaceae are generally canopy or emergent trees, with alternate, simple leaves, and trunks with fibrous bark (embira or envira). Some species have economic value, including Bertholletia excelsa Bonpl. (Brazil nut), used for food (seeds); Couroupita guianensis Aubl. (abricó- de-macaco), an ornamental valued for its flowers and cannonball-like fruits growing from the trunk; and Cariniana spp. (jequitibás) and Couratari spp. (tauaris), used for timber (Mori et al., 2014; Prance & Mori, 1979; Procópio & Secco, 2008). The family has a pantropical distribution, encompassing 17 genera and about 300 species, with the main center of diversity in the Guianan and Amazonian regions (Prance & Mori, 2004). In Brazil there are 10 genera and 119 species, most of them occurring in the Amazon domain (Smith et al., 2014). The Atlantic Forest, with 20 species of which 60% are endemic, is a secondary center of diversity of the family in Brazil (Mori, 1995; Smith et al., 2014). The members of Lecythidaceae are indicators of undisturbed or little- disturbed lowland forests (Mori et al., 2007); however, some species such as Eschweilera nana (O. Berg) Miers are small trees found in savanna or secondary habitats, where they are exposed to fire (Prance, 1975; Mori, 1990). No specific taxonomic treatments of the family in Espírito Santo have been published. However, studies such as the Flora Neotropica (Prance & Mori, 1979; Mori & Prance, 1990) and the synopsis of the species of eastern Brazil (Mori, 1995) have indicated that there are seven and nine species of Lecythidaceae in Espírito Santo, respectively. Mori (1995) emphasized that incomplete collections hinder the resolution of taxonomic problems of some taxa in this region. The List of Species of the Brazilian Flora records four genera and 10 species of the group for the state (Smith et al., 2014). 66 RIBEIRO ET AL.: LECYTHIDACEAE IN ESPÍRITO SANTO, BRAZIL Biological collections, as well as the systematic research associated with them, are invaluable for the knowledge, exploration and history of biodiversity. Specimens and publications provide the essential information needed to recognize areas with poorly known biota and to determine which areas should have priority for research and conservation of understudied taxonomic groups (Marinoni & Peixoto, 2010). Development of protocols for the conservation and sustainable use of plants, based on research and practical experience, and assessing the conservation status of plants are important, because knowledge of the species in a given region constitutes the first goal of the Global Strategy for Plant Conservation (GSPC, 2010), i.e., understanding and documenting plant diversity. The goal of this study was to inventory the Lecythidaceae occurring in Espírito Santo, gather information about previous collections, and determine areas and environments where species of Lecythidaceae occur. These steps will enable future studies that will support the goals of the GSPC. Material and Methods Information was initially obtained from the SpeciesLink network (CRIA, 2014), to determine which herbaria hold specimens of Lecythidaceae from Espírito Santo (BOTU, CEPEC, CVRD, ESA, FUEL, HPL, INPA, MBM, MBML, MO, NY, R, RB, SPF and UEC). Subsequently, we examined the specimens deposited in the herbaria of CVRD, MBML, RB, and VIES. The acronyms are listed following Thiers (2014). Identifications of specimens were confirmed or corrected when necessary. We also conducted fieldwork in 19 locations in Espírito Santo (Table 1), in which the