Moss diversity in the tropical rainforests of Rio de Janeiro, southeastern Brazil

Diversidade de musgos nas florestas tropicais do Rio de Janeiro, Sudeste do Brasil

Moss diversity in the tropical rainforests of Rio de Janeiro, southeastern Brazil

Diversidade de musgos nas florestas tropicais do Rio de Janeiro, Sudeste do Brasil

Denise P. da Costa¹; Fernanda M. Lima

Instituto de Pesquisas Jardim Botânico do Rio de Janeiro, Rua Pacheco Leão 915, 22460-030 Rio de Janeiro, RJ, Brasil

ABSTRACT

Moss diversity at various sites in the Tropical Atlantic Rainforest of southeastern Brazil is high, with 338 taxa distributed among 49 families and 129 genera. Comparisons of species richness in the Tropical Atlantic Rainforest in southeastern Brazil suggest that the moss flora is not uniform, and that lowland, montane, submontane, and upper montane Atlantic rainforests have very different moss floras. Montane Atlantic Rainforest has the largest number of exclusive species and the highest species richness, Sub-Montane Atlantic Rainforest has intermediate species richness, while the Lowland Atlantic Rainforest has fewer species. The high diversity of the Montane Atlantic Rainforest could be explained by the diversity of climatic, edaphic, and physiographic changes of the vegetation. Sematophyllaceae accounted for 19% of the taxa in lowland forest, Meteoriaceae for 10% of the taxa in montane forests, and Dicranaceae for 18% of the taxa in upper montane forests. Taxa with broad Neotropical distributions (40% of the total taxa) are important elements in all the forests, while taxa restricted to Brazil comprise the second most important element in upper montane and montane forests.

Key words: moss diversity, rainforests, southeastern Brazil

RESUMO

A diversidade de musgos na Floresta Tropical Atlântica do Sudeste do Brasil é considerada alta, com 338 táxons, distribuídos em 123 gêneros e 49 famílias. Comparações em relação à riqueza de espécies na Floresta Atlântica do Sudeste do Brasil sugerem que a flora de musgos não é uniforme e que as florestas das terras baixas, submontana, montana e altomontana apresentam floras muito diferentes. A Floresta Atlântica Montana conta com maior riqueza específica e número de táxons exclusivos, a Floresta Submontana tem riqueza intermediária e a Floresta Ombrófila Densa das Terras Baixas apresenta menor número de táxons. A alta diversidade da Floresta Montana pode ser explicada pelos fatores climáticos, edáficos e da vegetação. Sematophyllaceae conta com 19% dos táxons na floresta de terra baixa, Meteoriaceae com 10% dos táxons na floresta montana e Dicranaceae com 18% dos táxons na floresta altomontana. Os táxons neotropicais (40% do total de táxons) são elementos importantes para todas as florestas, enquanto que aqueles restritos ao Brasil são o segundo em importância para as florestas altomontana e montana.

Palavras-chave: diversidade de musgos, florestas tropicais, Sudeste do Brasil

Introduction

The Tropical Atlantic Rainforest of Brazil extends from the State of Rio Grande do Norte to Rio Grande do Sul in the coastal regions and inland on the mountains and plateau, with high humidity and rainfall. It has been suffering an intensive process of destruction by clear-cutting, shifting cultivation and human occupation, and only 1%-6% of the original area (1.2 million km²) persist in a mosaic of isolated fragments (Leitão Filho 1993). It is the most important ecosystem of Rio de Janeiro State (Fundação SOS Mata Atlântica 2002), and it has a rich moss flora.

In the southeast region of Brazil the forest extends further inland, where mountains can rise to 2,787 m in Itatiaia (Rio de Janeiro State) and 2,890 m on the Pico da Bandeira (Minas Gerais State). The original vegetation in this region was tropical rain forest, with small enclaves of Araucaria forest at higher altitudes and shrubby vegetation in the lower inland areas. The soils associated with eastern Atlantic Forest are mainly yellowish-redlatosols, clayey in texture. They occur in the half-orange mountains as well as in the watersheds of the main hydrographic basins. There are also yellowish-red clayey podzols occurring along the valleys in areas having a more pronounced dry season (Fundação SOS Mata Atlântica 2002, Leitão Filho 1993).

Regional endemism is high in the Tropical Atlantic Rainforest areas, around 55% for arboreal species and 40% for non-arboreal species (Barros et al. 1991, Joly et al. 1991, Mori et al. 1981, Peixoto & Gentry 1990).

Most of the bryophytes of the Tropical Atlantic Rainforest are epiphytes and some species grow on living leaves. According to Frahm & Gradstein (1991), bryophytes may be useful tools for the construction of a general scheme of the altitudinal zonation in tropical rainforests, because they are excellent climate indicators, they comprise rather few species (which facilitates identification), they have wide geographical ranges, and they are a characteristic structural component of the tropical rainforests.

According to Gradstein (1995), the bryophyte diversity in the Tropical Atlantic Rainforest varies considerably with altitude, both in structure and in floristic composition. This paper is the first to examine the moss diversity in the Tropical Atlantic Rainforest in southeastern of Brazil, and compares the region's lowland, submontane, montane, and upper montane moss floras. The approach is similar to that of studies carried out in other tropical rainforests (Gradstein & Salazar-Allen 1992). Some recent floristic studies have been done by the first author with the mosses of the Atlantic Rainforest in Rio de Janeiro State (Costa 1995, 1997, 1999, Costa & Yano 1995).

Material and methods

The taxa are from four different sites in the Atlantic Rainforest of Rio de Janeiro State: Poço das Antas (Silva Jardim), APA-Cairuçu (Parati), Nova Friburgo, and Itatiaia. Floristic inventories were conducted randomly by the first author at the first three sites between 1986-1993, and the data for Itatiaia are from the literature and herbarium records.

The vegetation classification applied to Atlantic Rainforest is that of Veloso et al. (1991), where Lowland Atlantic Rainforest = 0-50 m; Submontane Atlantic Rainforest = 50-500 m; Montane Atlantic Rainforest = 5001,500 m; and Upper Montane Atlantic Rainforest > 1,500 m. All the taxa were separated per altitudinal intervals according to classification of Veloso et al. (1991), being diversity and composition analysed per interval.

Floristic similarities between forest formations were calculated by means of the Sørensen Index of similarity. To estimate the real number of taxa per altitudinal interval from the observed number of taxa, the nonparametric S_chao estimate was used (Chao 1984). Chao's estimator is S_chao= S_obs+ F₁²/2F₂, where S_obs is the number of observed taxa, F₁ = the number of taxa with one record, and F₂ = the number of taxa with two records (Colwell 1997).

The classification for mosses is that of Buck & Goffinet (2000). The nomenclature follows Crosby et al. (1999).

All the samples collected are deposited at the herbarium RB, with duplicates to other herbaria on exchange. Study areas in Rio de Janeiro State – Poço das Antas is an important wildlife refuge of Lowland Atlantic Rainforest, comprising about 5,000 ha, 40% of it degraded and in distinct successional stages. The elevation ranges from sea level to 205 m and defines a vast plain subjected to phreatic or river flooding. Annual local rainfall averages approximately 1,600 mm annually. Rounded hills emerge from these floodplains, the highest ones reaching 205 m. Mean temperature is 22 ºC, ranging from 8 ºC to a maximum of 3032 ºC (IBDF 1981). Bryophyte cover is poor, the species are inconspicuous and mostly restricted to the canopy. Eighty six specimens of 37 taxa in 27 genera and 16 families were collected during three field trips carried out in 1994 (Costa 1999).

The "Área de Proteção Ambiental de Cairuçu" (Parati) is an important protected area of Submontane and Montane Atlantic Rainforest in the "Serra do Mar" range in southern Rio de Janeiro State, comprising about 33,800 ha. The elevation varies between 800-1,200 m above sea level, with the highest peak reaching 2,000 m. Air humidity averages 80%. Annual local rainfall averages 1,500-2,000 mm. Mean temperature is 22 ºC, ranging from 8 ºC to a maximum of 38 ºC (Conti et al. 1987). Bryophytes are abundant on tree trunks and branches. Two hundred and twenty specimens of 81 taxa in 44 genera and 20 families were collected during four field trips carried out between 1990 and 1991 (Costa 1997).

Nova Friburgo is an area of about 93,300 ha in the Montane and Upper Montane Atlantic Rainforest, in the "Serra dos Órgãos" range. Elevation varies between 8502,000 m above sea level, with the peak reaching 2,200 m. Air humidity averages 83%. Local rainfall averages 1,500 mm. Mean temperature is 25 ºC, ranging from 9 ºC to a maximum of 27 ºC (Brasil 1970). Bryophytic vegetation is luxuriant, and the tree trunks, branches and terrestrial layer are covered with a dense layer of bryophytes, including many robust and pendant species (Meteoriaceae, Pterobryaceae). A total of 514 specimens of 152 taxa in 77 genera and 34 families were collected during twelve fields trips carried out in 1986-1990 (Costa 1995).

Itatiaia is the first national park of Brazil, and an area of the Montane and Upper Montane Atlantic Rainforest in northwestern Rio de Janeiro State, in the "Serra da Mantiqueira" range. It comprises about 30,000 ha, 22º19'-22º45' S and 44º45'-44º50' W. Elevation varies between 6002,787 m above sea level, with the highest peaks reaching 2,408 m (Cabeça do Leão), 2,500 m (Dois Irmãos), and 2,787 m (Pico do Itatiaiaçu). Air humidity averages 85%. Local rainfall averages 2,100 mm (Hueck 1972). Mean temperature is 11 ºC, ranging from 6 ºC to a maximum of 27 ºC (Brade 1956, Hueck 1972, Pádua & Coimbra Filho 1979). The aspect of the bryophyte vegetation is similar to that of Nova Friburgo. Floristic data for Itatiaia are from Dusén (1903), Frahm (1991), Müller (1898), Reese (1993), Schäfer-Verwimp (1992), Schäfer-Verwimp & Giacontti (1993), Schäfer-Verwimp & Vital (1989), Yano (1992), and from the RB herbarium. A total of 273 samples were studied to Itatiaia. In total, 203 taxa in 95 genera and 42 families of mosses are found in Itatiaia.

Specific questions addresses here are: Does the species richness increase along an altitudinal gradient? Does the number of exclusive taxa increase along an altitudinal gradient? Does the number of endemic taxa increase along an altitudinal gradient? Can the moss flora be used to characterize the elevation belts in the Atlantic rainforest of southeastern Brazil? Could be some species considered indicators for different tropical Atlantic rainforests in southeastern Brazil?

Results and Discussion

Total diversity – In total, 338 taxa of mosses in 123 genera, and 49 families have been identified from the four study areas. We recorded, per altitudinal intervals, 68 taxa in Lowland Atlantic Rainforest, 90 taxa in Submontane Atlantic Rainforest, 202 taxa in Montane Atlantic Rainforest, and 161 in Upper Montane Atlantic Rainforest (table 1). The generic diversity is also high (123), with 55% (68) being represented by only one taxa. The differences observed in the floristic composition in the tropical rainforests of Rio de Janeiro seem to be related to habitat heterogeneity due primarily to topographic relief, which is in agreement with the conclusion of Churchill (1991). According to Churchill (1991), in the Neotropics, high diversity is related to habitat heterogeneity coupled with vegetational zonation. For this reason moss species diversity is related with the dramatic topographic relief found in this region (Churchill et al. 1995).

Floristics – In terms of number of taxa, Dicranaceae (45 taxa), Pilotrichaceae and Sphagnaceae (24 taxa), Sematophyllaceae (22 taxa), and Meteoriaceae (21 taxa), account for 40% (136 taxa) of the total taxa (table 1). Six taxa (9%) are exclusive to Lowland forest, and 9 taxa (10%) to Submontane forest, whereas 102 taxa (50%) were found only in the Montane forest, and 90 taxa (56%) are exclusive to Upper Montane forest (figure 1). The Montane and Upper Montane forests have the largest number of exclusive species. Similar results were obtained by Gradstein & Salazar-Allen (1992) in a montane rainforest in Panama.

Many families found in the Upper Montane forest are lacking in the Lowland forest (e.g. Andreaeaceae, Bruchiaceae, Catagoniaceae, Ditrichaceae, Entodontaceae, Hypopterygiaceae, Miniaceae, Phyllogoniaceae, Polytrichaceae), in the Montane forest (e.g. Andreaeaceae, Splachnobryaceae and Symphyodontaceae), and in the Submontane forest (e.g. Andreaeaceae, Bruchiaceae, Catagoniaceae, Ditrichaceae, Entodontaceae, Ephemeraceae, Funariaceae, Grimmiaceae, Mielichoferiaceae, Mniaceae, Rigodiaceae, Seligeriaceae, Sphagnaceae, Splachnaceae, and Symphyodontaceae).

Sematophyllaceae is the largest family in the Lowland and Submontane forests, fifth in importance to Montane and Upper Montane forests. Dicranaceae is the largest family in the Montane and Upper Montane forests. Meteoriaceae is an important family to Montane forest but its importance decreases significantly in the Lowland forest because this family has predominantly pendent species that require lower temperature, higher light levels, and higher air humidity not found in the lowland (Richards 1984).

Some genera have their highest diversity in Submontane forest (e.g. Calymperes, Callicostella), in Montane forest (e.g. Bryum, Campylopus, Leucobryum) or in the Upper Montane forest (e.g. Breutelia, Bryum, Campylopus, Fissidens, Schlotheimia, Sphagnum).

Nine taxa were common to four sites (figure 1): Chrysohypnum diminutivum, Fissidens zollingeri, Neckeropsis undulata, Pilosium chlorophyllum, Pilotrichella flexilis, Plagiothecium novogranatense, Sematophyllum subsimplex, S. swartzii, and Thuidium recognitum (2.6% of the total taxa). Meanwhile, 30 taxa were common to three sites (8.9% of the total taxa) and 97 taxa were common to two sites (28.6% of the total taxa).

General distribution patterns – Analysis of the species distributions showed ten distribution patterns: widespread (57 taxa), pantropical (17 taxa), Holarctic (1 taxa), Southern Hemisphere (1 taxa), Afro-America (7 taxa), tropical and subtropical America (27 taxa), neotropical (125 taxa), southern South America (4 taxa), disjunct: Neotropics-India, Northern and Southern hemisphere, and Brazil-India (3 taxa), and endemic to Brazil (95 taxa). Taxa with broad neotropical distributions comprised about 37.1% of the total, those restricted to Brazil about 28.2%, and widespread about 16.9% (table 2), which is similar with the results found by other authors for tropical rainforests (Gradstein & Salazar-Allen 1992, Gradstein et al. 1989).

In the lowland Atlantic rainforest, characteristic Neotropical taxa (35%) include Groutiella apiculata, and tropical and subtropical American taxa (13%) include Thamniopsis incurva. In the submontane Atlantic rainforest, Neotropical taxa (46%) include Calymperes lonchophyllum, and the widespread taxa (17%) include Meteorium nigrescens. In the montane Atlantic rainforest, Neotropical taxa (40%) include Bryohumbertia filifolia, widespread taxa (17%) include Hedwigidium integrifolium, and taxa restricted to Brazil (23%) include Mittenothamnium submacrodontium. In the upper montane Atlantic rainforest, taxa restricted to Brazil (33%) include Breutelia grandis, while Neotropical taxa (32%) include Campylopus capitulatus, and widespread taxa (18%) include Sphagnum capilifolium. It thus appeared that in the lowland, submontane and montane floras, species are usually widespread in the Tropical America, while the upper montane forests contain more taxa with restricted distribution. This is in agreement with the pattern in the Colombian Andes (Gradstein et al. 1989). Species richness – The total number of species in each elevational interval supports the notion that tropical Upper Montane and Montane forests are more diverse than the Submontane and Lowland forests (Richards 1984). The differences observed in the moss taxa richness for the four sites analyzed probably are due to habitat heterogeneity coupled with the vegetational zonation provided by topographic relief (Churchill et al. 1995).

The highest diversity is found in Montane and Upper Montane forests (202 and 161 taxa, figure 1), and may be explained by the higher rainfall, humus-rich soils, greater topographic relief, constantly high air humidity, lower temperatures, and higher light intensities, providing a diversity of microclimates and microhabitats inside these forests.

The lower number of taxa in the Lowland Forest can be explained by the open structure, absence of topographic relief, higher temperatures, higher light intensities, that affect the structure and internal microclimate in the forest.

The intermediate taxa diversity found in the Submontane forest (90 taxa, figure 1), could be explained by the intermediary conditions of this forest.

The high taxa richness of the forests above 500 m supports the notion that the Tropical Montane and Upper Montane forests have more moss taxa than Lowland forest under some conditions (Churchill et al. 1995, Vitt 1991).

The richness of mosses in the four altitudinal intervals ranged from 41-93 to the genera and 68-202 to the taxa (table 3). The highest richness is found in the Montane (93 genera and 202 taxa) and Upper Montane forests (77 genera and 161 taxa), and the lower in the Lowland forest (41 genera and 68 taxa). The Submontane forest (50 genera and 90 taxa) has intermediary richness. Using the nonparametric S_chao estimate, to extrapolate the real number of taxa per altitudinal interval from the observed number of taxa, we found that the real number is almost the same the observed number in the Lowland and Submontane forests, although is underestimated in the montane and Upper Montane forests (table 4).

It appears that in the Atlantic Rainforest of Southeastern of Brazil, high moss diversity commences in the montane zone between 500-2,700 m. According to Churchill et al. (1995), Frahm & Gradstein (1991), and Gradstein (1995), the causes of the change with the elevation of epiphytic bryophytes in the tropical rain forests are incompletely understood, and altitudinal related climatic factors such as the frequency of fog, high air humidity, air temperature or light intensity in the forest, or combinations of these factors, have been considered important.

Taxa × vegetational formations – Some taxa are here considered useful for the construction of a general scheme of the altitudinal zonation in the Tropical Rainforests in Rio de Janeiro, in another words, a characteristic structural component of the different vegetational formations along the elevational gradient (mainly based on the elevation ranges and geographical distributions).

Lowland – Acroporium exiguum and Homaliodendron piniforme.

Submontane – Campylopus trachyblepharum, Fissidens radicans, Isodrepanium lentulum, and Leucomium strumosum.

Montane – Campylopus dichrostis, Cryphaea malmei, Cryptopapillaria pennicilata, Cyclodictyon limbatum, Diphyscium ulei, Hookeriopsis rubens, Leiomella piligera, Lepidopilum plebejum, L. stenodictyon, Neckera caldensis, Oligotrichum riedelianum, Orthostichella microcarapa, O. mucronatula, Philophyllum tenuifolium, Pilotrichella nudiramulosa, Toloxis imponderosa, Sphagnum costae, S. rotundatum, S. rotundifolium, S. subovalifolium, and S. sucrei.

Upper Montane – Andreaea microphylla, A. spurioalpina, A. squarrosofiliformis, Aptychopsis pyrrophylla, Atractylocarpus brasiliensis, Brachydontium notorogenes, Brachymenium hornschuchianum, Breutelia grandis, B. subtomentosa, Campylopus fuscocroceus, C. julicaulis, Cladostomum ulei, Dicranella gymna, Ditrichum liliputianum, D. ulei, Ephemarum pachyneurum, Fissidens wallissii, Itatiella ulei, Macromitrium eriomitrium, Mielichhoferia grammocarpa, M. serrae, M. stridens, Neckera araucarieti, Orthodontium itacolumitis, O. pelluscens, Pohlia crassicostata, P. grammocarpa, Polytrichum angustifolium, P. brasiliense, Rhynchostegium rivale, Schlotheimia pseudoaffinis, S. trichomitria, Sematophyllum panduraefolium, Sphagnum gracilescens, S. minutulum, S. perforatum, S. platyphylloides, S. pseudoramulinum, Tetraplodon itatiaiae, Trachyxiphium drepanophyllum, Trematodon brevifolius, T. gymnostomus, and T. pauperifolius.

Excluded taxa – Here are included taxa not found in the Mosses World Checklist (Crosby et al. 1999) and in the literature or that are considered insufficiently known taxonomically: Bartamia rufescens Hampe (Itatiaia), Campylopus canalliculatus (Geh. & Hampe) Par. (Itatiaia), C. collinus Par. (Itatiaia), Eurhynchium riparioides (Hedw.) Richs. (Friburgo), Fissidens obtusatus Hampe (Itatiaia), F. longifalcatus Müll. Hall. (southeastern and southern Brazil), Mielichhoferia linearis Müll. Hall. (Itatiaia), Trematodon brevifolius Müll. Hall. (Itatiaia), T. gymnostomus Lindb. (Itatiaia), T. heterophyllus Müll. Hall. (Itatiaia), T. pauperifolius Müll. Hall. (Itatiaia).

Acknowledgments – The authors are grateful to Dr. Rogério Oliveira (Pontifícia Universidade Católica) and Dr. Douglas C. Daly (The New York Botanical Garden) for making valuable suggestions in the manuscript. Finally, the authors want to thank Dr. R.K. Colwell (Department of Ecology and Evolutionary Biology, University of Connecticut) for helping with the EstimateS program.

(received: December 18, 2003; accepted: October 13, 2005)

• BARROS, F., MELO, M.M.R.F., CHIEA, S.A.C., KIRIZAWA, M., WANDERLEY, M.G.L. & JUNG-MENDAÇOLLI, S.L. 1991. Caracterização geral da vegetação e listagem das espécies ocorrentes. In Flora Fanerogâmica da Ilha do Cardoso (M.M.R.F. Melo, F. Barros, M.G.L. Wanderley, M. Kirizawa, S.L. Jung-Mendaçolli & S.A.C. Chiea, eds.). Instituto de Botânica, São Paulo, p.1-184.
• BRADE, A.C. 1956. A Flora do Parque Nacional do Itatiaia. Jornal do Comércio, Rio de Janeiro.
• BRASIL. 1970. Observações Meteriológicas - Mod. D.M.A.- 1.110, período de 1931 a 1970 Departamento Nacional de Metereologia, Rio e Janeiro.
• BUCK, W.R. & GOFFINET, B. 2000. Morphology and classification of mosses. In Bryophyte biology (A.J. Shaw & B. Goffinet, eds.). Cambridge University Press, Cambridge, p.71-123.
• CHAO, A. 1984. Nonparametric estimation of the number of classes in a population. Scandinavian Journal of Statistics 11:265-270.
• CHURCHILL, S.P. 1991. The floristic composition and elevational distribution of Colombian Mosses. The Bryologist 94:157-167.
• CHURCHILL, S.P., GRIFFIN III, D. & LEWIS, M. 1995. Moss diversity of the tropical Andes. In Biodiversity and conservation of Neotropical Montane Forests (S.P. Churchill H. Balslev, E. Forero & J.L. Luteyn, eds.). The New York Botanical Garden, New York, p.335-346.
• COLWELL, R.K. 1997. EstimateS: Statistical estimation of species richness and shared species from samples. Version 5. User's Guide and application published at: http://viceroy.eeb.uconn.edu/estimates
• CONTI, V.M., MACIEL, N.C., MAURY, C.M.R.F. & COSTA, M.L.M.N. 1987. APA-Cairuçu-Parati-RJ: informações básicas. Sema - Secretaria do Meio Ambiente, Brasília.
• COSTA, D.P. 1995. Musgos do Município de Nova Friburgo, Rio de Janeiro, Brasil. Dissertação de mestrado, Universidade de São Paulo, São Paulo.
• COSTA, D.P. 1997. Bryophyta, Hepatophyta. In Mapeamento da cobertura vegetal e listagem das espécies ocorrentes na Área de Proteção Ambiental de Cairuçu, Município de Parati, RJ (M.C.M. Marques, coord.). Série Estudos e Contribuições 13:37-43.
• COSTA, D.P. 1999. Epiphytic bryophyte diversity in primary and secondary lowland rainforests in southeastern Brazil. The Bryologist 102:320-326.
• COSTA, D.P. & YANO, O. 1995. Musgos do Município de Nova Friburgo, Rio de Janeiro, Brasil. Arquivos do Jardim Botânico do Rio de Janeiro 33:99-118.
• CROSBY, M.R., MAGILL, R.E., ALLEN, B. & HE, S. 1999. A checklist of the mosses. Missouri Botanical Garden, Saint Louis.
• DUSÉN, P. 1903. Sur la flore de la Serra do Itatiaia. Arquivos do Museu Nacional do Rio de Janeiro 13:1-119.
• FRAHM, J.-P. 1991. Dicranaceae: Campylopodioideae, Paraleucobryoideae. Flora Neotropica. Monograph 54:1-238.
• FRAHM, J.-P. & GRADSTEIN, S.R. 1991. An altitudinal zonation of tropical rain forests using bryophytes. Journal of Biogeography 18:669-678.
• FUNDAÇÃO SOS MATA ATLÂNTICA. 2002. Atlas da evolução dos remanescentes florestais e ecossistemas associados do domínio da Mata Atlântica no período 1995-2000. Fundação SOS Mata Atlântica/INPE, São Paulo.
• GRADSTEIN, S.R. 1995. Bryophyte diversity of the tropical rainforest. Archives des Siences Genève 48:91-96.
• GRADSTEIN, S.R. & SALAZAR-ALLEN, N. 1992. Bryophyte diversity along an altitudinal gradient in Dárien National Park, Panama. Tropical Bryology 5:61-71.
• GRADSTEIN, S.R., VAN RENNEN & GRIFFIN III, D. 1989. Species richness and origin of the bryophyte flora of the Colombian Andes. Acta Botanica Neerlandica 38:439-448.
• HUECK, K. 1972. As Florestas da América do Sul. Universidade de Brasília, Polígono, São Paulo.
• IBDF. 1981. Plano de Manejo. Reserva Biológica de Poço das Antas. Fundação Instituto Brasileiro de Geografia e Estatística, Brasília.
• JOLY, C.A., LEITÃO FILHO, H.F. & SILVA, S.M. 1991. O Patrimônio Florístico. In Mata Atlântica (I.G. Câmara, coord.). Index/Fundação SOS Mata Atlântica, São Paulo, p.97-125.
• LEITÃO FILHO, H.F. (ORG.) 1993. Ecologia da Mata Atlântica em Cubatão. Editoras Unesp/Unicamp, São Paulo/Campinas.
• MORI, S.A., BOOM, B.M. & PRANCE, G.T. 1981. Distribution patterns and conservation of eastern Brazilian coastal forest tree species. Brittonia 33:233-245.
• MÜLLER, C. 1898. Bryologia Serrae Itatiaiae. Bulletin de l'Herbier Boissier 6:18-48.
• PÁDUA, M.T.J. & COIMBRA FILHO, A.F. 1979. OS Parques Nacionais do Brasil. Instituto de Cooperação Iberoamericana Instituto de la Caza Fotográfica y Ciencias de la Natureza, Madrid.
• PEIXOTO, A.L. & GENTRY, A. 1990. Diversidade e composição florística da mata de tabuleiro na Reserva Florestal de Linhares (Espírito Santo, Brasil). Revista Brasileira de Botânica 13:19-25.
• REESE, W.D. 1993. Calymperaceae. Flora Neotropica Monograph 58:1-102.
• RICHARDS, P.W. 1984. The Ecology of Tropical Forest Bryophytes. In New Manual of Bryology (R.M. Schuster, ed.). The Hattori Botanical Laboratory, Nichinan, v.2, p.1233-1270.
• SCHÄFER-VERWIMP, A. 1992. New or interesting records of Brazilian bryophytes, III. Journal of the Hattori Botanical Laboratory 71:55-68.
• SCHÄFER-VERWIMP, A. & GIANCOTTI, C. 1993. New or interesting records of Brazilian bryophytes, IV. Hikobia 11:285-292.
• SCHÄFER-VERWIMP, A. & VITAL, D.M. 1989. New or interesting records of Brazilian bryophytes. Journal of the Hattori Botanical Laboratory 66:255-261
• VELOSO, H.P., RANGEL FILHO, A.L.R. & LIMA, J.C.A. 1991. Classificação da vegetação brasileira adaptada a um sistema universal. IBGE/CDDI, Departamento de Documentação e Biblioteca, Rio de Janeiro.
• VITT, D.H. 1991. Distribution patterns, adaptive strategies, and morphological changes of mosses along elevational and latitudinal gradients on South Pacific Islands. In Quantitative approaches to phytogeography (P.L. Nimits & T.J. Crovello, eds.). Kluwer Academic Publishers, Leiden, p.205-231.
• YANO, O. 1992. Leucobryaceae (Bryopsida) do Brasil. Tese de doutorado, Universidade de São Paulo, São Paulo.

Publication Dates

• Publication in this collection
  01 Aug 2006
• Date of issue
  Dec 2005

History

• Received
  18 Dec 2003
• Accepted
  13 Oct 2005