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Reproductive Phenology, Pollination and Seed Dispersal Syndromes on Sandstone Outcrop Vegetation in the “Chapada Diamantina”, Northeastern Brazil: Population and Community

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Reproductive Phenology, Pollination and Seed Dispersal Syndromes on Sandstone Outcrop Vegetation in the “Chapada Diamantina”, Northeastern Brazil: Population and Community Revista Brasil. Bot., V.30, n.3, p.475-485, jul.-set. 2007 Reproductive phenology, pollination and seed dispersal syndromes on sandstone outcrop vegetation in the “Chapada Diamantina”, northeastern Brazil: population and community analyses1 ABEL AUGUSTO CONCEIÇÃO2,4, LIGIA SILVEIRA FUNCH2 and JOSÉ RUBENS PIRANI3 (received: February 23, 2006; accepted: August 23, 2007) ABSTRACT – (Reproductive phenology, pollination and seed dispersal syndromes on sandstone outcrop vegetation in the “Chapada Diamantina”, northeastern Brazil: population and community analyses). Vegetation on rock outcrops in the “Chapada Diamantina” (soil islands) is often aggregated and surrounded by nude rock surfaces, thus creating natural units with well defined limits. The flowering and fruiting cycles of plants on 58 soil islands at altitudes between 1,100 and 1,140 meters above sea leavel were analyzed at Mãe Inácia Peak (12°27’ S and 41°28’ W) in the “Chapada Diamantina”, Bahia, Brazil. The presence/ absence of flowering and fruiting species on each soil island, and their respective cover areas were analyzed at both the population and community levels, and the phenophases of flowering and fruiting were observed during 24 successive months. The analyses of pollination and seed dispersal syndromes indicated that animals are more important in pollination than in seed dispersal (which is predominantly by anemochory and autochory). The flowering and fruiting of plants with animal pollination syndromes were correlated with rainfall and temperature. The flowering season varied during the year according to the pollination syndrome involved: entomophily was predominant from summer through autumn, ornithophily was predominant during winter, and anemophily in the spring. The staggered timing of flowering and fruiting among different species provides a nearly continuous supply of resources for the local fauna. Key words - flowering, fruiting, insular sandstone outcrop vegetation, phenology, syndromes RESUMO – (Fenologia reprodutiva e síndromes de polinização e dispersão de semente em vegetação sobre afloramentos de arenito na Chapada Diamantina, Nordeste do Brasil: análises de população e comunidade). Parte da vegetação sobre afloramentos rochosos na Chapada Diamantina são agregados limitados pela superfície rochosa sem plantas vasculares (ilhas de solo), constituindo unidades naturais com limites bem definidos. Os ciclos de floração e frutificação de 58 ilhas de solo foram analisados no Morro da Mãe Inácia (12°27’ S e 41°28’ W), em altitudes entre 1.100 e 1.140 metros acima do nível do mar, na Chapada Diamantina, Bahia, Brasil. As presenças e ausências de espécies com flores e frutos em cada ilha de solo e respectivas áreas de cobertura foram analisadas nos níveis de população e comunidade. As fenofases de floração e frutificação nas ilhas foram realizadas durante 24 observações mensais. As análises das síndromes de polinização e dispersão de sementes indicaram que os animais são mais importantes para polinização do que para dispersão (predominantemente anemocoria e autocoria). A floração e frutificação das plantas que apresentam síndromes de polinização por animais foram correlacionadas com precipitação e temperatura. O período de floração variou de acordo com as síndromes de polinização envolvidas: entomofilia predominou do verão ao outono, ornitofilia predominou durante o inverno e anemofilia durante a primavera. Os períodos intercalados de floração e frutificação das espécies garantem contínuo suprimento de recursos para fauna local. Palavras-chave - fenologia, floração, frutificação, síndromes, vegetação insular em afloramento de arenito Introduction available to many organisms. Pollination and dispersal are crucial to plant reproduction, and to their ability to The phenological characteristics of plants are establish new populations on unoccupied sites. It is a essential elements in vegetation community dynamics, critical component of the maintenance of biological and they exercise a strong influence on resources diversity (Monasterio & Sarmiento 1976, Primack & Miao 1992, Newstrom et al. 1994, Howe & Westley 1997). Plant communities on small, well-defined areas on 1. This work is part of a PhD thesis developed at the Universidade de São Paulo, Departamento de Botânica, São Paulo, SP, rock outcrops are limited to species that can tolerate Brazil. (or avoid) severe restrictions of water, nutrients, and 2. Universidade Estadual de Feira de Santana, Departamento de soil (Collins et al. 1989, Burrows 1990, Ware 1990). Ciências Biológicas, BR 116, km 03, 44031-460 Feira de Santana, BA, Brazil. These restrictions are accentuated on mountain tops, 3. Universidade de São Paulo, Instituto de Biociências, where marked oscillations of wind speed and temperature Departamento de Botânica, Caixa Postal 11461, 05422-970 make establishment and growth even more difficult São Paulo, SP, Brazil. 4. Corresponding author: [email protected] (Steyermark 1986, Safford & Martinelli 2000). These 476 A. A. Conceição et al.: Phenology and syndromes on sandstone outcrop vegetation special communities completely surrounded by rock of daily minimums below 4 °C (Nimer 1989). Daily temperature surfaces are called “soil mat communities” in Africa amplitudes vary on average from 8 to 10 °C during the day, (Hambler 1964), “island communities” in the United with the largest amplitudes occurring at the highest altitudes States (Burbanck & Platt 1964, McCormick et al. 1974, (Jesus et al. 1983). During the study period (2000 to 2002), Shure & Ragsdale 1977), and “soil islands” in Brazil there were four months with less than 20 mm of rainfall, while the months of February and November had the warmest (Meirelles 1996). average temperatures (figure 1C). Some of the highest peaks in northeastern Brazil The “campo rupestre” vegetation growing on Mãe Inácia are found in the Chapada Diamantina Mountain Range. Peak is composed of a mosaic of distinct communities Many endemic species are found in “campo rupestre” distributed heterogeneously across the landscape according vegetation occurring on mountain tops with large areas to the type of substrate. The vegetation on the rocky surfaces of exposed rock (Giulietti & Pirani 1988, Harley 1995, at the very top of the mountain is composed of herbaceous Giulietti et al. 1997, Conceição 2000). These rock outcrop plants, shrubs, and small trees, and is dominated by Velloziaceae, communities are dominated by a restricted group of Clusiaceae, Cyperaceae, Orchidaceae, and Melastomataceae. plants, including species of Velloziaceae (Conceição & The first family is responsible for approximately 44% of the Giulietti 2002, Conceição 2003, Conceição & Pirani 2005, total vegetation cover (Conceição & Pirani 2005). Soil islands are insular units composed of two or more Conceição et al. 2007) that can survive dehydration species of vascular plants sharing the same patch of soil that during dry periods but quickly re-hydrate with the first is otherwise surrounded by a bare rock surface devoid of rains (Gaff 1977, Meguro et al. 1977, Meirelles et al. 1997). any vascular plants (Conceição & Pirani 2005). The study The only phenological study of soil islands undertaken area was selected based on its high proportion of rock surfaces, in Brazil (Meirelles 1996) reported that flowering as well as its accessibility and reduced human interference. occurred mainly during the rainy season and that life- Two plots were established: a 100 x 320 m rectangle (32,000 form was an important factor related to the phenological m²), and a 100 x 100 m square (10,000 m²), each of then divided patterns of these plants, especially in the case of geophytes, into 10 x 10 m square sub-plots. Six sub-plots were chosen by for which all above-ground parts are absent during the lottery, and all of the soil islands entirely or partially included driest months. The studies examining aspects of pollination within these six areas were identified and numbered, yielding a total of 58 individual soil islands. and floral biology have been undertaken in areas of “campo The ground coverage of the different plant species rupestre” usually focused a low number of species (Sazima composing each island were visually estimated based on the 1978a,b, Borba & Semir 2001, Madeira & Fernandes 1999, vertical projection of all aerial plants parts. The surface areas Borba et al. 2001, Carmo & Franceschinelli 2002). Only of the islands themselves were estimated by measuring their a sigle study considered more complex systems, sampling longest axis (l) and greatest width (w), then, based on the all mellitophilous plants and the Apoidea bees in a 1 ha general form of the island, either the area of an ellipse (¼ l.w.p) area of “campo rupestre” (Faria & Camargo 1996). or a rectangle (l.w) was calculated. Voucher specimens of all The present study sought to detect the patterns of species studied are housed at the herbarium SPF (“Universidade flowering and fruiting in insular plant communities de São Paulo”). growing on rocky surfaces at “Chapada Diamantina”, as Phenology – The presence or absence of plants with flowers or fruits in each of the 58 soil islands studied was recorded well as to relate these patterns to climatic factors (rainfall on a monthly basis from November 2000 to October 2002, and temperature) and pollination and seed dispersal generating a binary matrix of their phenophases. Due to the syndromes. difficulty in distinguishing single individuals, especially among monocot species, clumps of each species were Material
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