Vascular Flora of the Babitonga Bay Region (Santa Catarina, Brazil): Diversity and Origins Der Naturwissenschaftlichen Fakultät der Friedrich-Alexander-Universität Erlangen-Nürnberg zur Erlangung des Doktorgrades vorgelegt von Jotham Ziffer Berger aus Tel Aviv Als Dissertation genehmigt von der Naturwissenschaftlichen Fakultät der Universität Erlangen-Nürnberg Tag der mündlichen Prüfung: 15.7.2008 Vorsitzender der Promotionskommission: Prof. Dr. Eberhard Bänsch Erstberichterstatter: Prof. Dr. Werner Nezadal Zweitberichterstatter: Prof. Dr. Karin Esemann de Quadros ii לסבתי רות ברגר iii CONTENTS ACKNOWLEDGEMENTS 4 1 INTRODUCTION 5 1.1 THE ATLANTIC FOREST 5 1.2 SCOPE OF THE SURVEY 8 1.3 AIMS AND OBJECTIVES 10 2 THE STUDY AREA 11 2.1 GEOGRAPHIC LOCALIZATION 11 2.2 CLIMATE 14 2.3 GEOLOGICAL FEATURES 19 2.4 VEGETATION 26 2.5 USE OF NATURAL RESOURCES 35 3 METHODOLOGY 37 3.1 SELECTION OF SAMPLING LOCATIONS 37 3.2 FIELD WORK 38 3.3 SURVEY OF HERBARIUM SPECIMENS AND PREVIOUS CHECKLISTS 39 3.4 DETERMINATION OF PLANT SPECIMENS 39 3.5 BIOGEOGRAPHIC SURVEY 41 3.6 APPLIED TERMINOLOGY 42 3.7 DATA ARRANGEMENT AND PROCESSING 45 4 RESULTS 48 DETAILED PLANT LIST 48 4.1 PTERIDOPHYTES 48 4.2 SEED PLANTS 57 1 4.3 SUMMARY STATISTICS 128 5 DISTRIBUTION PATTERNS AND ORIGINS OF THE FLORA 134 5.1 ENDEMISM 134 5.2 SPECIES OF THE ATLANTIC FOREST AND ADJACENT REGIONS 143 5.3 WIDESPREAD SPECIES OF TROPICAL AMERICA 146 6 DIVERSITY, DISTRIBUTION AND CONSERVATION OF SELECTED TAXONOMIC GROUPS 151 6.1 ASTERACEAE 151 6.2 POACEAE 151 6.3 CYPERACEAE 152 6.4 ORCHIDACEAE 153 6.5 LEGUMES 153 6.6 MYRTACEAE 155 6.7 BROMELIACEAE 155 6.8 FERNS 156 7 DIVERSITY AND DISTRIBUTION OF SELECTED ECOLOGICAL GROUPS 158 7.1 PLANTS OF MANGROVE FOREST , SALT MEADOWS , AND THEIR PERIPHERY 158 7.2 STRAND PLANTS 159 7.4 FOREST SPECIES 160 7.5 PLANTS OF TREE RESTINGAS 161 7.6 RUDERAL PLANTS 163 7.7 ALIEN SPECIES 164 7.8 INVASIVE SPECIES 164 7.9 ESCAPING SPECIES 165 7.10 RARE SPECIES 165 8 CONCLUSION 169 8.1 SUMMARY OF THE FINDINGS 169 8.2 MEASURES FOR THE CONSERVATION OF THE FLORA 170 2 8.3 FOR FUTURE RESEARCH 171 BIBLIOGRAPHY 174 SUMMARY 205 ZUSAMMENFASSUNG 207 LISTS OF TABLES AND ILLUSTRATIONS 209 APPENDIX I: MAPS OF DISTRIBUTION 211 LEBENSLAUF 215 3 ACKNOWLEDGEMENTS I am deeply indebted to my advisor Prof. Dr. Werner Nezadal for the help to realize this research and for the pleasant atmosphere in which I had the fortune to work. I am also grateful to Prof. Dr. Karin Esemann de Quadros for the help, counsel and, most importantly, kind friendship during the research period in Brazil. I would like to express my heartfelt gratitude to the staff in Univille and to its Rector, Paulo-Ivo Koehntopp. I owe my special thanks to Prof. Cláudio Tureck and his wife Simone for the tremendous, ceaseless support and care during my stay in Vila da Glória. My thanks are also due to Prof. Cynthia Hering Rinnert for the hospitality in the herbarium in Joinville and for helpful advice. This work would not be realizable without the thoughtfulness and consideration of the staff of the Museu Botânico Municipal in Curitiba. I acknowledge a special debt to Dr. Gerdt Hatschbach, Osmar dos Santos Ribas, Juarez Cordeiro, Eraldo Barboza, and Clarisse Poliquesi for the generous guidance, suggestions and valuable advice. I thank my dear friends Alon Kol for reviewing this dissertation and for the graphic edition, and Henry Siman Tov for the German translation of the summary. Finally I would like to thank Dr. Walter Welß, Ricarda Bartschat, Jacqueline Alfandari, Dr. Astrid Oppolzer, Rosane Schweidson, Uri Siman Tov and all the friends and colleagues for their precious support and encouragement. J.Z. Fürth, April 2008 4 1 INTRODUCTION 1.1 THE ATLANTIC FOREST THE ATLANTIC FOREST BIOME —A CENTER OF BIOLOGICAL DIVERSITY The Atlantic Forest is a moist tropical and subtropical biome extending along the Brazilian Atlantic coast from the coast of the state of Rio Grande do Norte southwards to the Uruguayan coast, also reaching the Argentinean Province of Misiones and wide parts of southern Paraguay (see figure 1.1). The northern portion of the biome is relatively narrow, bordering with the caatinga (dry shrubland); the middle portion runs alongside the cerrado (savanna); and the broadest southern portion borders on the pampas in the south and on the Gran Chaco semi-arid plains in the west. Figure 1.1: the original extent of the Atlantic Forest [source: Conservation International (2008)] The Atlantic Forest—or Mata Atlântica in Portuguese—is one of the world’s leading centers of endemism and biodiversity (Dean 1995), with approximately 8000 endemic plants and 650 vertebrate species (Myers et al. 2000). The biome has suffered severe exploitation and deforestation since Europeans first settled in South America, and lost approximately 92% of its original extent (see figures 1.1 and 1.2). Therefore, international conservation organizations recognize the Atlantic Forest as a biodiversity hotspot for conservation priorities (Myers et al. 2000, Galindo-Leal & Câmara 2003), and UNESCO declared it a Biosphere Reserve in 1993 (UNESCO 2008). 5 Figure 1.2: map of the southern part of the Atlantic Forest, indicating two large forest remnants: the coastal forests of Serra do Mar (Brazil), and the semi-deciduous inland forest of Misiones (Argentina); taken from Galindo-Leal & Câmara (2003) Figure 1.3: the interior of the Atlantic Forest, photographed in a hillside forest in Vila da Glória 6 PHYTOGEOGRAPHY OF THE SOUTHERN ATLANTIC FOREST : ONE BIOME – TWO PHYTOCHORIA The present monograph focuses on an Atlantic Forest remnant in the southern part of the Atlantic Forest. Phytogeographically, the area belongs to the South American floristic realm [after Cox (2001), comprising Takhtajan’s (1986) Neotropical Kingdon and the South American part of the Holoantarctic Kingdom], and to the Amazonian Domain (Cabrera & Willink 1973) or after Takhtajan (1986) to the extra-Amazonian Brazilian Region. The Atlantic Forest biome is divided into two phytochoria (i.e. floristic provinces): the Atlantic Province comprising the coastal forests, coastal scrubs and mangroves; and the Paraná Province comprising the inland Araucaria forests and the semi-deciduous forests of Paraná-Paraguay basin. Some characteristic species of the Paraná province are the Brazilian pine ( Araucaria angustifolia ), erva-mate ( Ilex paraguariensis ), Podocarpus lambertii and the tree fern Dicksonia sellowiana (Daly & Mitchell 2000, Takhtajan 1986). The Babitonga bay region, the area studied in this monograph, is situated in the southernmost part of the Atlantic Province . The Atlantic Province shows strong floristic affinity to the Amazonian forest, as both were linked during warm and humid periods via gallery forests (Daly & Mitchell 2000). They still share a significant number of species, for example Pera glabrata ,(Euphorbiaceae) and Pouteria durlandii (Sapotaceae). The Atlantic Forest became isolated from the Amazonian forest probably during late Terciary and allowed the evolution of new, unique species (Mori et al. 1981). The Atlantic Province is floristically distinguishable from the Amazonian forest by the high endemism and diversity rates of Myrtaceae and Bromeliaceae (Daly & Mitchell 2000) and by endemic Rubiaceae and Melastomataceae. BOTANICAL RESEARCH IN THE SOUTH -BRAZILIAN ATLANTIC FOREST The diversity of plants in the south-Brazilian coastal forests is the main topic of this monograph. Since late 1950s modern research on plant diversity has been conducted in southern Brazil, especially in the state of Santa Catarina. The foremost botanical work for the region is Raulino Reitz’s, and later Ademir Reis’ (1965- ) Flora Ilustrada Catarinense , including most vascular plant families in the area. New family treatments are continuing to appear. With considerable contribution by world-renowned taxonomists the flora covers a great deal of the plant inventory of southern Brazil with ecological and biogeographic data. The journal Sellowia from the Barbosa Rodrigues Herbarium in Itajaí (Santa Catarina) includes some additional family treatments for the state. For our study area and its proximity, botanists have made significant contributions. Veloso & Klein (1961) presented the first phytosociological survey of the vegetation around the Babitonga bay. In this survey they analyzed the species composition and the structures of plant associations of rain forests in the region. Though requiring a taxonomical update, this survey is a milestone in modern botanical research in southern Brazil. After quite a long interval—not until the 1990s—the Babitonga bay region returned to be systematically studied. The Volta Velha reserve in Itapoá served as a sampling location for several botanical studies. Negrelle (1995), Dorneles & Negrelle (2000), Salimon & Negrelle (2001), Boeger et al. (2004) and Vamosi (2006) among others treat the ecological aspects of the area with contributions to phytosociology, reproduction biology and regeneration processes; Behling & Negrelle (1997) analyzed the modern pollen rain in the reserve; Behling & Negrelle (2001) investigated pollen in sediments for paleoecological characterization of the region; and Labiak & Prado listed the epiphytic pteridophytes. The Volta Velha reserve is an 7 excellently studied locality for its accessibility, structure, good state of preservation, and for the tremendous friendliness of the Machado family who owns the reserve. Botanical investigations in the area of Vila da Glória, a locality situated in the heart of the Babitonga