Phytogeography of the liverwort flora of the Atlantic Forest of southeastern

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1 Journal of Bryology (2010) 32: 9 22 Phytogeography of the liverwort flora of the Atlantic Forest of southeastern Brazil NIVEA DIAS DOS SANTOS 1,2 and DENISE PINHEIRO DA COSTA 2 1 Escola Nacional de Botânica Tropical and 2 Instituto de Pesquisas Jardim Botânico do Rio de Janeiro, Brazil SUMMARY This work analyzes the phytogeographic patterns of the liverwort flora of the Atlantic Forest of the Rio de Janeiro State, south-eastern Brazil. The analysis was based on inventories made in fragments of Atlantic Forest in the state, collections from the RB herbarium, and information from the checklist and database of the bryoflora of Rio de Janeiro. The phytogeographic pattern for each taxon was characterized based on its current distribution. Three hundred and sixty liverwort taxa are recognized for Rio de Janeiro. The liverwort flora varies along an altitudinal gradient within the Atlantic Forest, with the montane belt having the greatest species richness (238), the highest number of exclusive taxa (63) and the greatest number of endemic species (23). The predominant phytogeographic element is of neotropical species (49%), followed by disjunct liverworts (18%). Forty-one taxa (11%) are endemic to the country, of which 34 are restricted to the Atlantic Forest. In the lowland and submontane formations species are wide-ranging, whereas the montane and upper montane formations are characterized by endemic species or those disjunct with the Andes. The liverwort flora emphasizes the importance of the fragments of Atlantic Forest in Rio de Janeiro as a centre of diversity and endemism, supporting 50% of the total liverwort species known in Brazil, 72% of those recorded from Atlantic Forest and 55% of the endemic species of the country. Species with an Afro-American and Andean disjunction make up a characteristic part of the liverwort flora, probably reflecting the effects of long-distance dispersal by air currents, migration over land before the breakup of the continents and climatic similarities between the high-altitude grasslands and the northern Andes. KEYWORDS: Brazil, Rio de Janeiro, altitudinal belts, bryophytes, Marchantiophyta, phytogeographic patterns, rain forest INTRODUCTION With an area of km 2, Brazil is the largest country of South America and occupies a significant part of the Neotropical zone as well as subtropical areas to the south of the country. Brazil presents an immense variety of vegetation types, such as forests, grasslands, savannas, steppes, restingas and mangroves (Veloso, Rangel Filho & Lima, 1991). Brazil is one of the 17 most species-diverse countries of the world containing an estimated 1.8 million animal and plant species (Lewinsohn & Prado, 2005). Its plant diversity is probably one of the richest with more than species, ca 19% of the global flora (Giulietti et al., 2005). The country has ca 1650 bryophyte species (Costa et al., 2007 unpublished; Gradstein & Costa, 2003), which corresponds to 11% of the global bryoflora and 41% of the Neotropical bryoflora (Gradstein, Churchill & Salazar- Allen, 2001). The Atlantic Forest, the oldest forest formation in the country, has been established for at least 70 million years (Leitão-Filho, 1987). It is distributed along 27u latitude extending from northeast (3uS) to south Brazil (30uS), including parts of Argentina and Paraguay (Silva & Casteleti, 2005), and has an altitudinal variation from sea level up to 2890 m elevation. The Atlantic Rainforest is currently isolated from other large forest formations (Amazon Forest and Andean forests) by the dry vegetation of the Brazilian Planalto (caatinga, cerrado and pantanal) and by the Chaco in the central depressions of South America (Silva & Casteleti, 2005). This geographic isolation has allowed the origin of unique biota, with a high level of endemism (Rizzini, 1997). Moreover, during its evolutionary history, there have occurred periods of contact with the biota of these other forests (Leitão-Filho, 1987; Rizzini, 1997; Fernandes, 2003). For this reason, it is believed that the Atlantic # British Bryological Society 2010 Received 12 June Revision accepted 20 October 2009 DOI: / X

2 10 N. D. SANTOS AND D. P. COSTA Forest has undergone several periods of biological differentiation (Silva & Casteleti, 2005), having among its biota many ancient elements that became differentiated during the Miocene and more recent elements that colonized the region during the Quaternary (Prance, 1982; Rizzini, 1997). These unique characteristics make the Atlantic Forest one of the most singular biogeographic regions of South America (Müller, 1973; Silva & Casteleti, 2005). It is estimated that this region supports from 1 to 8% of global biodiversity, with species of plants, of which 40% are endemic. Thus, the Atlantic Forest can be considered one of the 34 global hotspots, making it a priority for conservation (Myers et al., 2000; Mittermeier et al., 2004). However, about 93% of this forest has been lost and the remaining fragments continue to deteriorate due to the cutting of timber, illegal extraction of plants and animals, introduction of exotic species, uncontrolled urban expansion, industrialization and immigration (Galindo- Leal & Câmara, 2005). The bryoflora of the Atlantic Forest is considered highly diverse and for liverworts it is the richest region in Brazil (ca 130 genera, 500 species) and the third most important in the Neotropics after the northern Andes and Central America (Gradstein et al., 2001; Gradstein & Costa, 2003). In the State of Rio de Janeiro (hereafter referred to as Rio de Janeiro), Atlantic Forest fragments represent 19% of the territory (Fundação SOS Mata Atlântica, 2002), and still shelter a considerable diversity of bryophytes, with a total of 1036 taxa (698 mosses, 333 liverworts and 5 hornworts) recognized by Costa, Imbassahy & Silva (2005a). Rio de Janeiro is considered one of the centers of diversity and endemism in Brazil (Guedes-Bruni & Lima, 1997; Lima et al., 1997; Rocha et al., 2003). Due to its location and immense variety of geographic formations (beaches, restingas, mangrove swamps, lakes, forests, and high-altitude grasslands) that stretch from the mountains to the sea, the state is characterized by a significant biological diversity and high level of endemism, which demonstrates the importance of its biota (Projeto Flora do Estado do Rio de Janeiro, unpublished data). This work contributes to existing knowledge of the liverwort diversity of Atlantic Forest by means of an analysis of phytogeographic patterns of the taxa occurring in Rio de Janeiro. The concept of the Atlantic Forest (sensu lato) adopted by this study is that of Oliveira-Filho & Fontes (2000) which encompasses evergreen and semi-deciduous forests. Study area MATERIALS AND METHODS Rio de Janeiro, situated on the Atlantic coast of southeastern Brazil (20u289 23u329S and 40u579 44u539W), possesses an area of km 2 (Fundação CIDE, 2006) and includes within its territory different vegetational formations of the Atlantic Forest biome, such as dense evergreen forest, seasonally semi-deciduous forest and mixed evergreen forest, in addition to restinga and mangrove swamps (Ministério das Minas e Energia, 1983). Eight climatic regions ranging from super-humid to semiarid have been recognized. In general, the climate is tropical and defined by two seasons: rainy (summer) and dry (winter). The annual precipitation varies between 500 and 2500 mm. The highest humidity is found along the southern coast and in mountainous regions of the state, while the lowest is found along the north coast and in the depressions of the Rio Paraíba do Sul valley. In the highest zones of the Serra do Mar and Serra da Mantiqueira, the characteristic climate is super-humid mesothermic, without a dry season (Ministério das Minas e Energia, 1983; Marques & Martins, 1997). Montane and lowland regions are present within the region. The topography is rugged, the highest peaks include Pico das Agulhas Negras (2791 m), in Planalto de Itatiaia; Pedra dos Três Picos (2310 m), in Nova Friburgo municipality; Pedra do Sino (2263 m) and Pedra do Açu (2230 m) in Serra dos Órgãos, Teresópolis; and Pico da Macela (1840 m), in Paraty municipality (Fundação CIDE, 2006; Guedes-Bruni & Lima, 1997). In the portions above 2000 m elevation and on some peaks are found the high-altitude grasslands. These are a series of humid formations dominated by grassy vegetation, restricted to the highest mountainous regions of southeastern Brazil and have a high index of endemism (Martinelli & Bandeira, 1989; Safford, 1999a). In Rio de Janeiro, the two principal high-altitude grasslands are found in the planalto of Itatiaia (22u259S, 44u409W) and in the Serra dos Órgãos (22u289S, 43u029W); while those that occur on the Pico do Desengano (21u549S, 41u549W), Pico do Frade (22u139S, 42u049W) and Pico do Tinguá (22u359uS, 43u299W) are also noteworthy. Rocha et al. (2003) recognize five principal zones of forest fragments in the state, four of them found in the Serra do Mar and one in the Serra da Mantiqueira. These fragments have relatively continuous vegetation with some degree of connectivity and are referred to as the following: North Fluminense Zone (in the area of Serra do Desengano and Morro de São João), Central Serrana Zone (Três Picos, Serra dos Órgãos, Serra do Tinguá), Rio de Janeiro Metropolitan Zone (Maciço da Tijuca, Maciço da Pedra Branca and of the Serra do Mendanha-Gericinó), South Fluminense Zone (Serra da Bocaina, in the region of Paraty, Baía de Ilha Grande) and Serra da Mantiqueira Zone (in the region of Resende and Itatiaia). Liverwort survey Inventories were conducted in forest fragments in different altitudinal belts: lowland forests Poço das Antas Biological Reserve, Tijuca National Park (TNP), Desengano State Park (DSP); submontane TNP, DSP, El Nagual Private Reserve of Natural Heritage; montane Itatiaia

3 PHYTOGEOGRAPHY OF LIVERWORTS IN ATLANTIC FOREST 11 National Park (INP), Serra dos Órgãos National Park (SONP), TNP, DSP, APA Petrópolis; and upper-montane INP, SONP (all collections deposited in RB). Information was also obtained from collections in the herbarium of the Jardim Botânico do Rio de Janeiro (RB), from the Rio de Janeiro checklist (Costa et al., 2005a) and the Rio de Janeiro Bryoflora Database (Costa et al., 2007). The resulting list includes all species of liverworts known for the Atlantic Forest of Rio de Janeiro (Table 1). Specimens were identified by Gradstein & Costa (2003) and Fulford (1963, 1966, 1968, 1976), by means of comparison with herbarium material, and the help of specialists. The classification adopted for the Division Marchantiophyta is that proposed by Crandall-Stotler & Stotler (2000). Analyses For the phytogeographic analysis taxa were allocated to altitudinal zones according to the classification of vegetation by Veloso et al. (1991), where lowland forest55 to Table 1. Phytogeographic patterns of liverwort species in the altitudinal belts of the Atlantic Forest in Rio de Janeiro state, Brazil. *5endemic species of Rio de Janeiro state. For definition of phytogeographical patterns see Table 2. Altitudinal belt in Atlantic Forest (m) Liverwort Phytogeographical pattern Acanthocoleus aberrans (Lindenb. & Gottsche) Kruijt Af-Am X X Acrolejeunea emergens (Mitt.) Steph. Pan X X Acrolejeunea torulosa (Lehm. & Lindenb.) Schiffn. Neo X Adelanthus carabayensis (Mont.) Grolle Neo X Adelanthus decipiens (Hook.) Mitt. Wide X Anastrophyllum auritum (Lehm.) Steph. Pan X X Anastrophyllum piligerum (Nees) Steph. Pan X X Anastrophyllum tubulosum (Nees) Grolle Neo X X Aneura pinguis (L.) Dumort. Wide X X Anomoclada portoricensis (Hampe & Gottsche) Váña Neo X Anoplolejeunea conferta (Meissn.) A. Evans Neo X X X Aphanolejeunea asperrima Steph. ST X X Aphanolejeunea camillii (Lehm.) R.M. Schust. Neo X Aphanolejeunea clavatopapillata (Steph.) E. Reiner Other X Aphanolejeunea gracilis Jovet-Ast Neo X Aphanolejeunea microscopica var. africana (Pócs) Pócs & Bernecker Af-Am X Aphanolejeunea paucifolia (Spruce) E. Reiner Other X X Aphanolejeunea sintenisii (Steph.) Steph. Disj X X Aphanolejeunea truncatifolia Horik. Pan X X X Archilejeunea auberiana (Mont.) A. Evans Neo X X Archilejeunea fuscescens (Hampe ex. Lehm.) Fulford Neo X X Archilejeunea parviflora (Nees) Schiffn. Neo X X Aureolejeunea fulva R.M. Schust. Neo X Balantiopsis brasiliensis Steph. Atl X X Bazzania aurescens Spruce Neo X Bazzania cuneistipula (Gottsche & Lindenb.) Trevis. Disj X Bazzania heterostipa (Steph.) Fulford Atl X X Bazzania hookeri (Lindenb.) Trevis. Neo X X Bazzania jamaicensis (Lehm. & Lindenb.) Trevis. Neo X Bazzania longistipula (Lindenb.) Trevis. Neo X X X Bazzania nitida (Weber) Grolle Wide X Bazzania pallide-virens (Steph.) Fulford Neo X Bazzania schlimiana (Gottsche) Fulford Neo X X Bazzania stolonifera (Sw.) Trevis. Neo X Bazzania taleana (Gottsche) Fulford Neo X Blepharolejeunea incongrua (Lindenb. & Gottsche) Van Slageren & Kruijt Neo X Blepharolejeunea securifolia (Steph.) R.M. Schust. Neo X Brachiolejeunea laxifolia (Taylor) Schiffn. Neo X Brachiolejeunea phyllorhiza (Nees) Kruijt & Gradst. Af-Am X Bromeliophila natans (Steph.) R.M. Schust. Atl X Bryopteris diffusa (Sw.) Nees Neo X X X Bryopteris filicina (Sw.) Nees Neo X X X X Calypogeia grandistipula (Steph.) Steph. Atl X X Calypogeia laxa Gottsche & Lindenb. Neo X X X Calypogeia lechleri (Steph.) Steph. Neo X Calypogeia miquelii Mont. Neo X Calypogeia peruviana Nees & Mont. Neo X X

4 12 N. D. SANTOS AND D. P. COSTA Table 1. Continued. Altitudinal belt in Atlantic Forest (m) Liverwort Phytogeographical pattern Calypogeia uncinulatula Herzog Neo X X Caudalejeunea lehmanniana (Gottsche) A. Evans Af-Am X Cephalozia crassifolia (Lindenb. & Gottsche) Fulford Neo X X Cephalozia crossii Spruce Neo X Cephaloziella divaricata (Sm.) Schiffn. Other X X Cephaloziella granatensis (J.B. Jack) Fulford Disj X Cephaloziopsis intertexta (Gottsche) R.M. Schust. Neo X Ceratolejeunea ceratantha (Nees & Mont.) Steph. Disj X Ceratolejeunea cornuta (Lindenb.) Schiffn. Neo X X Ceratolejeunea cubensis (Mont.) Schiffn. Neo X X X Ceratolejeunea fallax (Lehm. & Lindenb.) Bonner Neo X X X Ceratolejeunea laetefusca (Austin) R.M. Schust. Neo X Ceratolejeunea rubiginosa Gottsche Neo X X X Cheilolejeunea acutangula (Nees) Grolle Neo X Cheilolejeunea clausa (Nees & Mont.) R.M. Schust. Neo X X X Cheilolejeunea discoidea (Lehm. & Lindenb.) Kachr. & R.M. Schust. Neo X X Cheilolejeunea holostipa (Spruce) R.-L. Zhu & Grolle Neo X X Cheilolejeunea inflexa (Hampe) Grolle Neo X X Cheilolejeunea insecta Grolle & Gradst. Other X Cheilolejeunea oncophylla (Ångstr.) Grolle & E. Reiner Neo X X X Cheilolejeunea rigidula (Mont.) R.M. Schust. Af-Am X X X Cheilolejeunea trifaria (Reinw. et al.) Mizut. Pan X X X Chonecolea doellingeri (Nees) Grolle Pan X X Clasmatocolea vermicularis (Lehm.) Grolle Other X X Cololejeunea cardiocarpa (Mont.) A. Evans Pan X X Cololejeunea minutissima (Sm.) Schiffn. Pan X X Cololejeunea minutissima ssp. myriocarpa (Sm.) Schiffn. Pan X X Cololejeunea obliqua (Nees & Mont.) Schiffn. Neo X X Cololejeunea subcardiocarpa Tixier Neo X X Cololejeunea vervimpii Tixier Neo X Colura calyptrifolia (Hook.) Dumort. Wide X *Colura itatyana Steph. Atl X Colura tenuicornis (A. Evans) Steph. Pan X X Colura ulei Jovet-Ast Neo X Cryptochila grandiflora (Lindenb. & Gottsche) Grolle Wide X *Cylindrocolea brasiliensis D.P. Costa & N.D. Santos Atl X Cylindrocolea planifolia (Steph.) R.M. Schust. Neo X Cylindrocolea rhizantha (Mont.) R.M. Schust. Neo X X X Dicranolejeunea axilaris (Nees & Mont.) Schiffn. Neo X Diplasiolejeunea alata Jovet-Ast Neo X X Diplasiolejeunea brunnea Steph. Neo X X Diplasiolejeunea cavifolia Steph. Pan X Diplasiolejeunea pauckertii (Nees) Steph. And X Diplasiolejeunea pellucida (Meissn.) Schiffn. Neo X X Diplasiolejeunea replicata (Spruce) Steph. And X Diplasiolejeunea rudolphiana Steph. Pan X Diplasiolejeunea unidentata (Lehm. & Lindenb.) Schiffn. Neo X X Drepanolejeunea anoplantha (Spruce) Steph. Other X X Drepanolejeunea araucariae Steph. AST X Drepanolejeunea bidens (Steph.) A. Evans Neo X Drepanolejeunea biocellata A. Evans Neo X Drepanolejeunea campanulata (Spruce) Steph. And X Drepanolejeunea fragilis Bischl. Neo X Drepanolejeunea granatensis (J.B. Jack & Steph.) Bischl. And X Drepanolejeunea inchoata (Meissn.) Schiffn. Neo X Drepanolejeunea lichenicola (Spruce) Steph. Neo X Drepanolejeunea mosenii (Steph.) Bischl. Neo X X X X Drepanolejeunea orthophylla (Nees & Mont.) Bischl. Neo X Drepanolejeunea palmifolia (Nees) Steph. Bra X Dumortiera hirsuta (Sw.) Nees Other X X X Fossombronia porphyrorhiza (Nees) Prosk. Neo X X X Frullania apiculata (Reinw. et al.) Nees Pan X Frullania arecae (Spreng.) Gottsche Pan X X X X

5 PHYTOGEOGRAPHY OF LIVERWORTS IN ATLANTIC FOREST 13 Table 1. Continued. Altitudinal belt in Atlantic Forest (m) Liverwort Phytogeographical pattern Frullania atrata (Sw.) Nees Neo X X Frullania beyrichiana (Lehm. & Lindenb.) Lehm. & Lindenb. Neo X X X Frullania brasiliensis Raddi Neo X X X X Frullania caulisequa (Nees) Nees Neo X X X X Frullania cuencensis Taylor Neo X Frullania dusenii Steph. Neo X X X X Frullania ecklonii (Spreng.) Gottsche et al. Pan X Frullania ericoides (Nees) Mont. Pan X X X X Frullania gaudichaudii (Nees & Mont.) Nees & Mont. Disj X Frullania gibbosa Nees Neo X X Frullania glomerata (Lehm. & Lindenb.) Mont. Neo X X Frullania intumescens (Lehm. & Lindenb.) Lehm. & Lindenb. Neo X Frullania kunzei (Lehm. & Lindenb.) Lehm. & Lindenb. Neo X X X X Frullania montagnei Gottsche Neo X Frullania mucronata (Lehm. & Lindenb.) Lehm. & Lindenb. Neo X X Frullania riojaneirensis (Raddi) Ångstr. Pan X X X X Frullania schaefer-verwimpii Yuzawa & S. Hatt. Atl X Frullania setigera Steph. Neo X Frullania supradecomposita (Lehm. & Lindenb.) Lehm. & Lindenb. Bra X X X Frullania vitalii Yuzawa & S. Hatt. Atl X Frullanoides corticalis (Lehm. & Lindenb.) Van Slageren Neo X X Frullanoides densifolia Raddi Neo X X Gongylanthus liebmanianus (Lehm & Lindenb.) Steph. Neo X Harpalejeunea oxyphylla (Nees & Mont.) Steph. Neo X X Harpalejeunea schiffneri S.W. Arnell Bra X X X Harpalejeunea stricta (Lindenb. & Gottsche) Steph. Neo X X Harpalejeunea subacuta A. Evans Disj X Herbertus grossispinus (Steph.) Fulford Neo X Herbertus juniperoideus ssp. acanthelius (Spruce) Feldberg & Heinrichs CAA X Herbertus juniperoideus ssp. bivittatus (Spruce) Feldberg & Heinrichs Neo X X Herbertus juniperoideus ssp. juniperoideus (Sw.) Grolle Neo X X Herbertus juniperoideus ssp. pensilis (Taylor) Spruce Neo X Herbertus oblongifolius (Steph.) Gradst. & Cleef And X Heteroscyphus combinatus (Nees) Schiffn. Neo X X Isotachis aubertii (Schwägr.) Mitt. Af-Am X X X *Isotachis inflata Steph. Atl X X Isotachis multiceps (Lindenb. & Gottsche) Gottsche Neo X X X Isotachis serrulata (Sw.) Gottsche Neo X Jamesoniella rubricaulis (Nees) Grolle Disj X Jensenia spinosa (Gottsche) Grolle Af-Am X X Jungermannia amoena Lindenb. & Gottsche Neo X X Jungermannia hyalina Lyell Holo X Jungermannia sphaerocarpa Hook. Holo X Kurzia brasiliensis (Steph.) Grolle Bra X X X Kurzia capillaris (Sw.) Grolle Af-Am X X X Kurzia flagellifera (Steph.) Grolle Neo X Kymatocalyx dominicensis (Spruce) Váña Disj X X X Lejeunea bermudiana (A.Evans) R.M. Schust. Disj X Lejeunea cancellata Nees & Mont. Neo X Lejeunea capensis Gottsche Pan X Lejeunea caulicalyx (Steph.) E. Reiner & Goda Neo X X X Lejeunea cerina (Lehm. & Lindenb.) Gottsche Neo X X Lejeunea controversa Gottsche Neo X X X Lejeunea cristulaeflora (Gottsche ex Steph.) E. Reiner & Goda Atl X Lejeunea cristulata (Steph.) E. Reiner & Goda Atl X X X Lejeunea flava (Sw.) Nees Pan X X X X Lejeunea grossiretis (Steph.) E. Reiner & Goda Atl X X Lejeunea grossitexta (Steph.) E. Reiner & Goda Other X X Lejeunea immersa Spruce Neo X X X X Lejeunea inflexiloba J.B. Jack & Steph. And X Lejeunea laeta (Lehm. & Lindenb.) Gottsche Neo X X Lejeunea laetevirens Nees & Mont. Neo X X X Lejeunea magnoliae Lindenb. & Gottsche Neo X X X

6 14 N. D. SANTOS AND D. P. COSTA Table 1. Continued. Altitudinal belt in Atlantic Forest (m) Liverwort Phytogeographical pattern Lejeunea monimiae (Steph.) Steph. Atl X X X *Lejeunea oligoclada Spruce Atl X X Lejeunea phyllobola Nees & Mont. Neo X X X Lejeunea puiggariana Steph. Atl X X Lejeunea raddiana Lindenb. Neo X X X Lejeunea reflexistipula (Lehm. & Lindenb.) Gottsche Neo X Lejeunea setiloba Spruce Neo X X X Lejeunea tapajosensis Spruce Neo X Lejeunea trinitensis Lindenb. Neo X Lepidolejeunea eluta (Nees) R.M. Schust. Neo X Lepidolejeunea involuta (Gottsche) Grolle Neo X Lepidozia coilophylla Taylor Neo X Lepidozia cupressina (Sw.) Lindenb. Wide X X Lepidozia inaequalis (Lehm. & Lindenb.) Lehm. & Lindenb. And X X Leptolejeunea brasiliensis Bischl. Disj X Leptolejeunea diversilobulata Bischl. Other X Leptolejeunea elliptica (Lehm. & Lindenb.) Schiffn. Neo X X X Leptolejeunea exocellata (Spruce) A. Evans Neo X Leptolejeunea moniliata Steph. Neo X X Leptoscyphus amphibolius (Nees) Grolle Neo X X Leptoscyphus gibbosus (Taylor) Mitt. Disj X Leptoscyphus porphyrius (Nees) Grolle Neo X Leptoscyphus spectabilis (Steph.) Grolle Atl X Lethocolea glossophylla (Spruce) Grolle CAA X Leucolejeunea conchifolia (A. Evans) A. Evans Disj X X Leucolejeunea unciloba (Lindenb.) A. Evans Af-Am X X X X Leucolejeunea xanthocarpa (Lehm. & Lindenb.) A. Evans Pan X X X X Lophocolea bidentata (L.) Dumort. Wide X X X Lophocolea connata (Sw.) Nees Neo X X Lophocolea glaziovii Steph. Atl X Lophocolea lindmannii Steph. Disj X Lophocolea mandonii Steph. Disj X X Lophocolea martiana Nees Af-Am X X X X Lophocolea martiana subsp. bidentula (Nees) Gradst. Atl X X Lophocolea muricata (Lehm.) Nees ST X X X Lophocolea perissodonta (Spruce) Steph. Neo X X Lophocolea quadridentata Spruce And X Lophocolea trapezoides Mont. AST X Lopholejeunea nigricans (Lindenb.) Schiffn. Pan X X X Lopholejeunea subfusca (Nees) Schiffn. Pan X X Lophozia bicrenata (Schmidel. ex Hoffm.) Dumort. Holo X Lunularia cruciata (L.) Dumort. Other X X Marchantia berteroana Lehm. & Lindenb. Other X Marchantia breviloba A. Evans Neo X Marchantia chenopoda L. Neo X X X Marchantia papillata Raddi AST X X X X Marchantia polymorpha L. Wide X Marchesinia brachiata (Sw.) Schiffn. Af-Am X X X Marsupella microphylla R.M. Schust. And X Mastigolejeunea auriculata (Wilson) Schiffn. Pan X X Mastigolejeunea plicatifolia (Spruce) Steph. Neo X Metalejeunea cucullata (Reinw. et al.) Grolle Pan X Metzgeria adscens Steph. Disj X Metzgeria agnewiae Kuwah. Pan X Metzgeria albinea var. albinea Spruce Af-Am X X X X Metzgeria albinea var. angusta (Steph.) Costa & Gradst. AST X Metzgeria aurantiaca Steph. Neo X X X X Metzgeria brasiliensis Schiffn. Atl X Metzgeria ciliata Raddi Other X X X X Metzgeria conjugata Lindb. Wide X X Metzgeria consanguinea Schiffn. Pan X Metzgeria convoluta Steph. Other X X Metzgeria cratoneura Schiffn. Atl X

7 PHYTOGEOGRAPHY OF LIVERWORTS IN ATLANTIC FOREST 15 Table 1. Continued. Altitudinal belt in Atlantic Forest (m) Liverwort Phytogeographical pattern Metzgeria dichotoma (Sw.) Nees Neo X X X Metzgeria fruticola Spruce Neo X Metzgeria furcata (L.) Dumort. Wide X X X Metzgeria herminieri Schiffner Neo X Metzgeria lechleri Steph. AST X X X Metzgeria leptoneura Spruce Wide X X Metzgeria myriopoda Lindb. Other X X X Metzgeria psilocraspeda Schiffn. Atl X X Metzgeria rufula Spruce Pan X Metzgeria scyphigera A. Evans AST X Metzgeria subaneura Schiffn. Bra X Microlejeunea bullata (Taylor) Steph. Neo X X X X Microlejeunea epihylla Bischl. Disj X Microlejeunea subulistipa Steph. Atl X Micropterygium pterygophyllum (Nees) Trevis. Other X Mnioloma cyclostipa (Spruce) R.M. Schust. Disj X X Monoclea gottschei subsp. elongata Gradst. & Mues Neo X X X X Myriocoleopsis gymnocolea (Spruce) E. Reiner & Gradst. Neo X Neesioscyphus argillaceus (Nees) Grolle And X X X Neesioscyphus carneus (Nees) Grolle Atl X X Neesioscyphus homophyllus (Nees) Grolle Atl X Neurolejeunea breutelii (Gottsche) A. Evans Neo X X X Noteroclada confluens Taylor ex Hook. & Wilson Af-Am X X Odontolejeunea decemdentata (Spruce) Steph. Neo X Odontolejeunea lunulata (Weber) Schiffn. Af-Am X Odontoschisma brasiliense Steph. Other X Odontoschisma denudatum (Nees) Dumort. Wide X X Odontoschisma falcifolium Steph. Other X X X Odontoschisma longiflorum (Taylor) Steph. Neo X Omphalanthus filiformis (Sw.) Nees Neo X X X Pallavicinia lyellii (Hook.) Gray Wide X X Paracromastigum dusenii (Steph.) R.M. Schust. Disj X Paracromastigum pachyrhizum (Nees) Fulford Neo X X Pictolejeunea picta (Gottsche ex Steph.) Grolle Neo X Pictolejeunea sprucei Grolle Bra X Plagiochasma rupestre (Forster) Steph. Other X Plagiochila adiantoides (Sw.) Lindenb. Neo X Plagiochila bifaria (Sw.) Lindenb. Other X X Plagiochila boryana Gottsche Af-Am X Plagiochila corrugata (Nees) Nees & Mont. Af-Am X X X X Plagiochila cristata (Sw.) Lindenb. Neo X Plagiochila disticha (Lehm. & Lindenb.) Lindenb. Neo X X Plagiochila distinctifolia Lindenb. Neo X X Plagiochila diversifolia Lindenb. & Gottsche Neo X Plagiochila exigua (Taylor) Taylor Wide X X Plagiochila flaccida Lindenb. Neo X X Plagiochila gymnocalycina (Lehm. & Lindenb.) Lindenb. Neo X X X Plagiochila macrostachya Lindenb. Neo X X Plagiochila martiana (Nees) Lindenb. Neo X X X Plagiochila micropteryx Gottsche Neo X X Plagiochila montagnei Nees Neo X X X Plagiochila patentissima Lindenb. Neo X X X Plagiochila patula (Sw.) Lindenb. Neo X X X Plagiochila raddiana Lindenb. Neo X X Plagiochila rutilans Lindenb. Neo X X X X Plagiochila simplex (Sw.) Lindenb. Neo X X X Plagiochila subplana Lindenb. Neo X X Pluvianthus squarrosus (Steph.) R.M. Schust. & Schäf.-Verw. Atl X Porella brasiliensis (Raddi) Schiffn. Atl X X X Porella reflexa (Lehm. & Lindenb.) Trevis. Atl X X X Prionolejeunea aemula (Gottsche) A. Evans. Neo X X X Prionolejeunea denticulata (Weber) Schiffn. Neo X Prionolejeunea limpida Herzog Atl X

8 16 N. D. SANTOS AND D. P. COSTA Table 1. Continued. Altitudinal belt in Atlantic Forest (m) Liverwort Phytogeographical pattern Pycnolejeunea densistipula (Lehm. & Lindenb.) Steph. Neo X Radula angulata Steph. Neo X Radula fendleri Gottsche Other X Radula gottscheana Taylor Neo X Radula javanica Gottsche Pan X X X X Radula kegelii Gottsche ex. Steph. Neo X X Radula ligula Steph. Atl X X Radula mexicana Lindenb. & Gottsche AST X X Radula nudicaulis Steph. Disj X X X Radula obovata Castle Other X X Radula pocsii K. Yamada Other X Radula quadrata Gottsche AST X Radula recubans Taylor AST X X X X Radula schaefer-verwimpii K. Yamada And X X Radula sinuata Gottsche Other X X Radula tectiloba Steph. AST X Radula tenera Mitt. CAA X Radula voluta Taylor ex Gottsche Wide X Rectolejeunea berteroana (Gottsche) A. Evans Neo X Riccardia amazonica (Spruce) S.W. Arnell Neo X Riccardia cataractatum (Spruce) Schiffn. AST X X X Riccardia chamedryfolia (With.) Grolle Holo X X X X Riccardia digitiloba (Spruce ex Steph.) Pagán Neo X X X X Riccardia emarginata (Steph.) Hell Atl X X X Riccardia fucoidea (Sw.) Schiffn. Neo X X X Riccardia glaziovii (Spruce) R. Meenks Neo X X X Riccardia metzgeriiformis (Steph.) Schiffn. Neo X X X Riccardia multifida (L.) Gray Holo X Riccardia regnellii (Ångstr.) Hell Bra X X X Riccia plano-biconvexa Steph. AST X X Riccia stenophylla Spruce Neo X X Riccia wainionis Steph. Neo X Saccogynidium caldense (Ångstr.) Grolle Atl X X Scapania portoricensis Hampe & Gottsche Neo X Schiffneriolejeunea polycarpa (Nees) Gradst. Pan X X X *Southbya organensis Herzog Atl X Stephaniella paraphyllina J.B. Jack Af-Am X Stictolejeunea squamata (Willd.) Schiffn. Neo X X X Symbiezidium barbiflorum (Lindenb. & Gottsche) A. Evans Neo X X Symbiezidium transversale (Sw.) Trevis. Neo X X Symphyogyna aspera Steph. Neo X X X X Symphyogyna brasiliensis (Nees) Nees & Mont. Af-Am X X X Symphyogyna brongniartii Mont. Neo X X Symphyogyna podophylla (Thunb.) Mont. & Nees ST X X X X Syzygiella anomala (Lindenb. & Gottsche) Steph. CAA X Syzygiella integerrima Steph. Neo X Syzygiella liberata Inoue CAA X Syzygiella perfoliata (Sw.) Spruce Neo X X Syzygiella uleana Steph. Atl X X Taxilejeunea isocalycina (Ness) Steph. Neo X Taxilejeunea lusoria (Lindenb. & Gottsche) Steph. Neo X X Taxilejeunea obtusangula (Spruce) A. Evans Neo X X Taxilejeunea pterigonia (Lehm. & Lindenb.) Schiffn. Neo X X X X Telaranea diacantha (Mont.) Engel & Merr. Pan X X X X Telaranea nematodes (Gottsche) M.A. Howe Af-Am X X X X Triandrophyllum subtrifidum (Hook.f. & Taylor) Fulford & Hatch. Other X Trichocolea brevifissa Steph. Neo X X Trichocolea flaccida (Spruce) J.B. Jack & Steph. Neo X Trichocolea tomentosa (Sw.) Grolle Neo X Tylimanthus laxus (Lehm. & Lindenb.) Spruce Neo X Vitalianthus bischlerianus (Pôrto & Grolle) R.M. Schust. & Giancotti Atl X

9 PHYTOGEOGRAPHY OF LIVERWORTS IN ATLANTIC FOREST m, submontane forest550 to 500 m, montane forest5500 to 1500 m, and upper-montane forest.1500 m. The phytogeographic patterns were based on global distributions of taxa, according to information contained in the Banco de Dados da Brioflora do Estado do Rio de Janeiro (Costa et al., 2007) and in the literature. The patterns were characterized following the geographical ranges of the taxa adapted from those described in the literature (Cabrera & Willink, 1980; Gradstein & Costa, 2003). RESULTS Diversity and floristic composition We identified 31 families, 102 genera, 354 species, 5 subspecies and 1 variety of liverworts, with 41 taxa endemic to Brazil (11%). Among these latter, 34 are restricted to the Atlantic Forest and 5 are endemic (Table 1). A total of 111 taxa occur in lowland forests with 19 restricted to this formation (5%); 167 taxa occur in the submontane forests, with 18 restricted (5%); 238 taxa are found in montane forests, with 63 restricted (17%); and 173 taxa are found in uppermontane forests, with 58 restricted taxa (16%). Thirty-one taxa (8.6%) occur in the four altitudinal belts, 65 (18%) in at least three belts, and 106 (29.4%) in at least two, with the hightest number of taxa (45) shared between the montane and upper-montane forests. The most well represented families are Lejeuneaceae (42 genera, 139 spp.), Lepidoziaceae (6 genera, 22 spp.), Jubulaceae (1 genus, 22 spp.), Plagiochilaceae, Metzgeriaceae (1 genus, 21 spp.), Geocalycaceae (5 genera, 17 spp.), and Radulaceae (1 genus, 17 spp.). The genera with the greatest number of species are Lejeunea (26 spp.), Frullania (22 spp.), Plagiochila, Metzgeria (21 spp. each), Radula (17 spp.), Drepanolejeunea (12 spp.), Bazzania (11 spp.), Lophocolea and Riccardia (10 spp. each). Some families, such as Acrobolbaceae, Arnelliaceae, Gymnomitriaceae and Scapaniaceae, are restricted to the upper-montane formations (.1500 m), while Adelanthaceae, Herbertaceae, Jungermanniaceae and Trichocoleaceae are restricted to montane and upper-montane formations (.500 m). Phytogeographic patterns Twelve phytogeographic patterns were identified (Tables 1, 2; Fig. 1) with the majority of taxa (49%) having a Neotropical distribution. Among those with disjunct patterns (18%), the Afro-American (19 taxa), Andean (11 taxa), combined Central America Andes Atlantic Forest of south-eastern Brazil (five taxa) and the Holarctic (five taxa) are noteworthy. Forty-one taxa are endemic to Brazil (11%), of which 34 are endemic to Atlantic Forest, with 29 restricted to the Brazilian Atlantic Forest and five extending to northern Argentina (Lejeunea monimiae, L. puiggariana, Porella brasiliensis, P. reflexa and Radula ligula), also distributed in the inland Atlantic Forest (Galindo-Leal & Câmara, 2005). Among the taxa endemic to Brazil (Fig. 2), the greatest proportions are confined to the montane (68%) and upper-montane (51%) forest. Pantropical taxa account for 8% of the liverwort flora and those with widespread global distributions amount to only 4%. Taxa with tropical and subtropical American distribution (TSA) make up 4% of the flora. Included in this latter group are Amphilejeunea reflexistipula, Drepanolejeunea araucariae, Lejeunea laeta, Lophocolea trapezoides, Marchantia papillata, Riccardia cataractatum and Riccia plano-biconvexa. Some species classified as Neotropical have a distribution restricted to South America: Bazzania pallide-virens, Mastigolejeunea plicatifolia, Myriocoleopsis gymnocolea, Riccardia glaziovii, R. metzgeriiformis and Riccia stenophylla. The species whose phytogeographic patterns could not be defined are included in Other, for example, Clasmatocolea vermicularis and Triandrophyllum subtrifidum, which occur in temperate regions of the Southern Hemisphere and in the tropical mountains of high elevation from the Andes to Costa Rica and south-eastern Brazil, the latter species also Table 2. Phytogeographical patterns of liverworts of Atlantic Forest in Rio de Janeiro state, with their acronyms and taxa numbers. Phytogeographic pattern Acronym Taxa number Widespread (found at least in three continents) Wide 15 Pantropical (wide tropical) Pan 30 Tropical and subtropical America TSA 12 Neotropical Neo 176 Holartic (disjunct between temperate regions of northern hemisphere and south-eastern Brazil) Holo 5 Tropical amphi-atlantic (disjunct between Africa and America) Af-Am 19 Andino Element (disjunct between northern Andes and Atlantic Forest of south-eastern of Brazil) And 11 Disjunct between the high mountains of Central America, Andes and south-eastern Brazil CAA 5 Southern temperate Element (disjunct between temperate regions of southern hemisphere and south-eastern Brazil) st 3 Other disjunctions Disj 19 Endemic of Brazil (not restricted to Atlantic Forest) Bra 7 Atlantic element (restricted to Atlantic Forest) Atl 34 Pattern not identified Other 24

10 18 N. D. SANTOS AND D. P. COSTA Figure 1. Table 2. Phytogeographical patterns of liverwort taxa of Rio de Janeiro, emphasizing disjunct patterns. For definition of acronyms see Figure 2. Phytogeographical patterns of liverwort taxa of the state of Rio de Janeiro in different altitudinal belts of the Atlantic Forest. For definition of acronyms see Table 2.

11 PHYTOGEOGRAPHY OF LIVERWORTS IN ATLANTIC FOREST 19 occurs in tropical Asia. Another, Plagiochasma rupestre, is distributed in tropical and Mediterranean regions. Distribution within altitudinal belts At all elevations in the forest the predominant phytogeographic pattern is Neotropical (Fig. 2). However, for the lowland and submontane forests, Pantropical species (18 and 11%, respectively) and Afro-American species (8 and 7%) predominate, whereas in the montane and uppermontane formations, species endemic to the Atlantic Forest (9 and 10%) and Pantropical species (8% in both formations) are most frequent. Taxa with disjunct distributions predominate in the upper-montane formation (44 taxa, 25%). Those with a disjunction between the Atlantic Forest and the high mountains of the Andes and/or Central America and the Andes (14 taxa) are noteworthy; these patterns are practically exclusive to this formation. In the lowland, submontane and montane formations, the prominent disjunction is Afro-American, with nine species occurring in the first, 11 in the second, and 15 in the third formation. Excluded species Twelve species were considered insufficiently well known taxonomically, were cited for the state only once, without locality information, have not been collected again, or have been included on the red list of the IUCN. Doubtful taxonomic status Lejeunea anomala Lindenb. & Gottsche, Lejeunea glauscescens Gottsche, Lejeunea lepida Lindenb. & Gottsche, Plagiochila lingua Steph. Locality unknown Bazzania gracilis (Hampe & Gottsche) Steph., Leptolejeunea maculata (Mitt.) Schiffn., Marchantia paleacea Bert., Radula stenocalyx Mont., Riccia curtisii (James) Austin, Riccia grandis Nees, Xylolejeunea crenata (Nees & Mont.) X.-L. He & Grolle. Included on the red list of the IUCN Drepanolejeunea aculeata Bischl. an epiphyll endemic to Atlantic Forest, known from only two localities in south-eastern Brazil (RJ, SP) and not encountered since 1922, being classified as endangered (EN) in the IUCN list. DISCUSSION Our results demonstrate the outstanding conservation importance of the Atlantic Forest remnants in Rio de Janeiro for liverworts. More than 50% of the liverwort species known for the whole country are recorded here and some 72% of the liverwort flora for the Brazilian Atlantic Forest as a whole (Gradstein & Costa, 2003; Santos, 2008). Of the 74 species of liverworts endemic to Brazil (Gradstein & Costa, 2003), 41 occur in Rio de Janeiro (55%), highlighting the importance of the Atlantic Forest of Rio de Janeiro as a center of biodiversity and endemism for the country. Among the 41 endemic species, seven occur in other vegetational formations in the country, predominantly the Amazon Forest (Gradstein & Costa, 2003), while 34 are restricted to Atlantic Forest. Among these species five are endemic to Rio de Janeiro including Colura itatyana, Isotachis inflata, Lejeunea oligoclada and Southbya organensis in montane and/or upper-montane formations, and Cylindrocolea brasiliensis in the submontane formation. Therefore, 7% of the species endemic in Brazil are restricted to the Rio de Janeiro forests, a high number given that about 12.5% (74 spp.) of the country s liverwort flora is considered endemic (Gradstein & Costa, 2003). The percentage of liverwort species endemic to Atlantic Forest is high when compared to other Neotropical countries. Additionally, 11 (31%) of the 35 families of liverworts that occur in the country are exclusive to or mainly recorded from this biome (Gradstein et al., 1994; Gradstein & Costa, 2003). As for the species of vascular plants, the degree of endemism is even greater for the Atlantic Forest, according to Guedes-Bruni & Lima (1997), the Rio de Janeiro forests have 70 80% endemics. According to Rizzini (1997) the high indices of biodiversity and endemism for Atlantic Forest could have originated during the Quaternary, when climatic fluctuations resulted in the processes of spatial expansion and retraction of the Atlantic Forest from more restricted forests that served as refuges for the flora and fauna. Today, these refuges are found in southern Bahia, in the region of Tabuleiros in Espírito Santo, along the coast of Rio de Janeiro and in the north of São Paulo. These regions exhibit high species diversity and endemism (Rizzini, 1997; Rambaldi et al., 2003). Floristic composition varies considerably with altitude within the Atlantic Forest. The montane belt has the highest species richness and the most endemic species, followed by the upper-montane belt, which has the greatest proportion of unique species. Similar results were found in an analysis of the mosses by Costa & Lima (2005). The high diversity of bryophytes in these formations can be explained by a variety of climatic, edaphic, and floristic factors including high levels of humidity and soil humus, and the open canopy which allows substantial illumination of the forest floor. These conditions favour the growth of species on the soil surface and on the trunk bases of trees (Gradstein, 1995). There are many microhabitats, which together with the steep relief makes a high species richness possible (Churchill, Griffin & Lewis, 1995). In addition to this, the montane belt represents a floristic transition zone as many genera have their lower elevational limits (Anastrophyllum, Herbertus, Jungermannia, Syzygiella, Trichocolea, etc.) or upper limits (Acanthocoleus, Archilejeunea, Ceratolejeunea, Cylindrocolea, Fossombronia, etc.) in this belt. Wolf (1993), studying the bryophytes of the Colombian montane forests, also found similar

12 20 N. D. SANTOS AND D. P. COSTA zonation patterns and affirmed that the transition zones generally exhibit high diversity as they are meeting points between distinct floristic assemblages. Our analysis demonstrates a change in liverwort phytogeographic elements as well as in floristic composition with increasing altitude of the Atlantic Forest. This phenomenon has been encountered in other tropical regions (e.g. Schuster, 1983; Gradstein, Van Reenen & Griffin, 1989; Gradstein, 1995). In the lowland and submontane areas Neotropical species predominate, together with Pantropical and Afro- American taxa which have a very wide distribution extending into the Paleotropics. In montane and uppermontane forest, above the Neotropical taxa, liverworts with a restricted distribution, including many endemics and those disjunct with populations in the northern Andes and in Central America and the Andes, are preponderant. A similar pattern was observed by Gradstein (1995) for the liverwort flora of the tropical Andean forests. Among the disjunct liverworts, of particular note is the Afro-American disjunction which Gradstein, Pócs & Váña (1983) regarded as a typical feature of the tropical liverworts. They hypothesized that this results from the long-distance dispersal of diaspores by transatlantic air currents. Such species generally produce small (,25 mm diam.) diaspores in great quantity. These are also resistant to desiccation and cold (Van Zanten & Pócs, 1981). Moreover, these species possess the ability to compete successfully with indigenous vegetation as many of them are found in areas of secondary vegetation (Gradstein et al., 1983). Examples are Isotachis aubertii, Lophocolea martiana and Symphyogyna brasiliensis. Van Zanten & Gradstein (1988) investigated the dispersal ability of Neotropical and transoceanic liverwort species experimentally and proposed a hypothesis of long-distance dispersal or land-based migration of spores for the disjunctions between Africa and tropical America. The latter could have involved stepby-step migration before the separation of the two continents from Gondwana. They verified that the tolerance of spores to desiccation is a principal factor determining success. Additionally, species which occur at altitudes below 800 m have a greater dispersal potential than those at higher elevations. It is possibly for these reasons that the Afro-American disjunction predominates in the liverwort flora of Rio de Janeiro s lowland and submontane Atlantic Forests. Some 16 hepatic species of upper-montane Atlantic Forest show a disjunction with the Andes, in some cases, reaching as far north as Central America. Six of these are restricted to high-altitude grasslands and/or transitional areas between forests and meadows: Drepanolejeunea granatensis, Lejeunea inflexiloba, Lethocolea glossophylla, Marsupella microphylla, Syzygiella anomala and S. liberata. Gradstein & Costa (2003) commented that there is a similarity between the high plateaux of south-eastern Brazil and the páramos, particularly in Itatiaia, and that evidence exists that this type of disjunction results from spore dispersal over long distances. Safford (2007), analyzing the vascular plant genera of the high-altitude grasslands of Brazil, also concluded that their floristic similarity is greater with the tropical Andes than with lowland and middle elevation areas of central and south-eastern Brazil. About 11% of the taxa of these grasslands are shared with the Andes. This may arise because of both current environmental similarities and shared biogeographical history (Safford, 1999b, 2007). The high-altitude grasslands represent a focus of Andean vegetation in Brazil that could disappear in the future with predicted climatic change (Safford, 2007). Palynofloras show that because the regional temperatures have warmed and become more humid, the high-altitude grasslands have significantly contracted over the past years, with a decrease of dominance of grassland and increase of forest taxa (Behling et al. 2007; Safford, 2007). The liverwort flora of these forests appears to show a similar phytogeographic make-up to the moss flora of Rio de Janeiro (Costa & Lima, 2005). Thus, the Neotropical element is especially prevalent for both groups. Overall, among Rio de Janeiro s mosses the Brazilian endemic (28%) and widespread (17%) elements predominate, while for liverworts the disjuncts (18%) are ranked first followed by the Brazilian endemics (11%). The moss flora also shows similar altitudinal patterns to the liverworts, the majority of endemic taxa occurring within the montane and uppermontane belts. There are similar proportions of endemic moss taxa in the upper-montane belt (33%) and among the Neotropical element (32%). The main differences between the moss and liverwort floras arise in the Itatiaia flora, which presents an exceptional number of endemic mosses and disjunct liverworts. These results corroborate the results obtained for Neotropical bryophytes (Gradstein & Costa, 2003; Costa, Imbassahy & Silva, 2005b), where the greatest number of endemic and disjunct species is found in the montane and upper montane zones. Studies of the vascular plant flora of Rio de Janeiro have reached even more remarkable conclusions. Lima et al. (1997) analyzed the flora of the Macaé de Cima Ecological Reserve (Nova Friburgo municipality), a region of Rio de Janeiro that has montane and upper-montane formations of Atlantic Forest. They found that 86.5% of the species are restricted to Brazil among which 91% are endemic to Atlantic Forest. Similar results were obtained for the arborescent Leguminosae (Lima, 2000) where the predominant element were the Brazilian endemics (56.8%). Much the same was true for the shrubby and arborescent Leguminosae of montane and upper-montane formations in the Itatiaia National Park (Morim, 2006), where Brazilian endemics amounted to 65% of the group. In comparison with these plants, the distributions of liverworts are wider and there are fewer endemics. In contrast to the flowering plants, the ferns show greater similarity to the bryophytes. Neotropical ferns were found to predominate in Restinga de Jurubatiba National Park (59%; Santos, Sylvestre & Araujo, 2004) and among the pteridophytes of

13 PHYTOGEOGRAPHY OF LIVERWORTS IN ATLANTIC FOREST 21 a coastal rock outcrop (37.5%; Santos & Sylvestre, 2006). Moreover, in the upper-montane region of the Itatiaia National Park, Condack (2006) found that Brazilian endemics were predominant (41.5%) and increased in proportion with increasing altitude as found here for liverworts and by Costa & Lima (2005) for mosses.the similarities between these groups are probably explained by the same features: the species are ancient, which has allowed time for a wider distribution to develop; the diaspores are produced in great quantity, and are small and resistant to desiccation, which favours their long range dispersal (Van Zanten & Pócs, 1981; Schofield, 2001). Contrary to what has been observed for vascular plants, endemic bryophytes are rare and taxa with transoceanic distributions are generally common, and for the most part, families and genera have a cosmopolitan distribution (Schofield, 1992). Overall, the results found for the liverwort flora of Atlantic Forest of Rio de Janeiro, where 11% of the flora is endemic, corroborate those observed for Atlantic Forest of south-eastern Brazil that is considered one of the centers of diversity and endemism in the Americas (Guedes-Bruni & Lima, 1997; Myers et al., 2000; Mittermeier et al., 2004; Pinto et al., 2006). ACKNOWLEDGEMENTS We thank Dr Maria Elena Reiner-Drehwald for help with identifications and in the determination of distribution patterns of some Lejeunea; the Fundação O Boticário de Proteção à Natureza (project no. 0709_20061) and the Instituto de Pesquisas Jardim Botânico do Rio de Janeiro for finance; the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) for a masters grant awarded to NDS; to Felipe W. Amorim for the critical reads to the early version of this manuscript; and to Jeff Bates and Steven P. Churchill for their helpful critiques and suggestions for improvements of final drafts. 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