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Survey of bryophytes in Serra da Canastra National Park, Minas Gerais, Brazil

ABSTRACT

Serra da Canastra National Park is located in southwestern Minas Gerais State and contains several phytophysiognomies of the Brazilian Cerrado. To date, no study on bryophytes has been conducted in this area and the present study is the first to be carried out in this major biological reserve of Cerrado located on the Brazilian Plateau. This study found 289 species of bryophytes, including mosses, liverworts, and one species of hornwort, representing fourty-one Brazilian endemic species and 56 new records for Minas Gerais State. Most species are widely distributed in Brazil, with only 16% having restricted distributions. Regarding worldwide distribution, 31% are Neotropical. Lejeuneaceae had the highest species richness among liverwort families with 53 species, whereas Sphagnaceae had the highest richness among moss families with 26 species. Phaeoceros laevis was the only hornwort species found in the park. This study contributes to the understanding of bryophyte species richness and distribution, and provides the worldwide and Brazilian distribution of the bryophyte flora of Serra da Canastra National Park, an important center for biodiversity conservation.

Keywords
cerrado; floristic; hornwort; liverworts; mosses; mountain areas

Introduction

Serra da Canastra National Park was created in order to protect the source of the São Francisco River, which is the watercourse arising in the huge chest-shaped plateau. Serra da Canastra divides two watersheds: the south portion is the beginning of the Paraná River and the north portion is the beginning of the São Francisco River (Souza & Costa 2011Souza A, Costa M. 2011. Parque Nacional da Serra da Canastra . http://www.serradacanastra.com.br/. 19 Aug. 2015.
http://www.serradacanastra.com.br...
).

The predominant vegetation is represented by grasslands and rocky fields (campos rupestres) (Carvalho-Silva & Guimarães 2009Carvalho-Silva M, Guimarães EF. 2009. Piperaceae do Parque Nacional da Serra da Canastra, Minas Gerais, Brasil. Boletim de Botânica da Universidade de São Paulo 27: 235-245.), which ensure a high degree of endemism since they occur in areas with particular ecological conditions influenced by geographic isolation and biotic and abiotic factors (Harley & Simmons 1986Harley RM, Simmons NA. 1986. Florula of Mucugê. Chapada Diamantina - Bahia, Brazil. Kew, Royal Botanical Garden.; Giulietti et al. 1987Giulietti AM, Menezes NL, Pirani JR, Meguro M, Wanderley MGL. 1987. Flora da Serra do Cipó, Minas Gerais: caracterização e lista das espécies. Boletim de Botânica da Universidade de São Paulo 9: 1-151.; Pirani et al. 1994Pirani JR, Giulietti AM, Mello-Silva R, Meguro M. 1994. Checklist and patterns of geographic distribution of the vegetation of Serra do Ambrósio, Minas Gerais, Brazil. Revista Brasileira de Botânica 17: 133-147.; Stannard 1995Stannard BL. 1995. Flora of the Pico das Almas, Chapada Diamantina - Bahia, Brazil. London, Kew, Royal Botanic Gardens.). Since the park is open to visitors there is some anthropic disturbance, such as the installation of unpaved roads and buildings; nevertheless, there is a relatively high degree of endemism among some angiosperm species, and so a great need for the permanent preservation of the area as a conservation unit (Romero & Nakajima 1999Romero R, Nakajima JN. 1999. Espécies endêmicas doParque Nacional da Serra da Canastra , Minas Gerais. Revista Brasileira de Botânica 22: 259-265).

Floristic surveys conducted in areas of Minas Gerais that are similar to Serra da Canastra include Yano & Carvalho (1995Yano O, Carvalho AB. 1995. Briófitas da Serra da Piedade, Minas Gerais, Brasil. In: Anais do 9o Congresso da Sociedade Botânica de São Paulo. São Paulo, Universidade Estadual Paulista. p.15-25.) who found 46 species in Serra da Piedade; Yano & Peralta (2009)Yano O, Peralta DF. 2009. Flora de Grão-Mogol, Minas Gerais. Briófitas (Bryophyta e Marchantiophyta). Boletim de Botânica da Universidade de São Paulo 27: 1-26. who found 42 species of mosses and hepatics in the Grão-Mogol mountains; Yano & Peralta (2011a)Yano O, Peralta DF. 2011a. Bryophytes from Serra de São José, Tiradentes, Minas Gerais, Brasil. Boletim de Botânica da Universidade de São Paulo 21: 141-172. found 114 species in Serra de São José at the city of Tiradentes; Yano & Peralta (2011b)Yano O, Peralta DF. 2011b. Flora da Serra do Cipó, Minas Gerais: Briófitas (Anthocerotophyta, Bryophyta e Marchantiophyta). Boletim de Botânica da Universidade de São Paulo 29: 135-211. found 237 species in Serra do Cipó; Luizi-Ponzo et al. (2013Luizi-Ponzo AP, Siviero TS, Amorim ET, et al 2013. Briófitas do Parque Estadual do Ibitipoca no Herbário Prof. Leopoldo Kriegeer. In: Forzza RC, Neto LM, Salimena FRG, Zappi D. (eds.) Flora do Parque Estadual do Ibitipoca e seu entorno. Vol. 4. 1st. edn. Juiz de Fora, Editora UFJF. p. 95-122.) found 209 species in Ibitipoca State Park and, recently, Souza & Câmara (2015Souza RV, Câmara PEAS. 2015. Survey of the bryophytes of a gallery forest in the National Park of Serra do Cipó, Minas Gerais, Brazil. Acta Botanica Brasilica 29: 24-29.) who studied the mosses of gallery forest in Serra do Cipó National Park. These works contain relevant floristic data and are useful in the identification of some species. Furthermore, they provide species lists and document the location of bryophytes in these regions, resulting in knowledge of about 766 species of bryophytes in Minas Gerais State (Costa & Peralta 2015Costa DP, Bastos CJP, Schäfer-Verwimp A. 2015. Lejeuneaceae In: Lista de Espécies da Flora do Brasil. Jardim Botânico do Rio de Janeiro. http://www.floradobrasil.jbrj.gov.br/jabot/floradobrasil/FB97355. 19 Aug. 2015.
http://www.floradobrasil.jbrj.gov.br/jab...
).

These works notwithstanding, floristic inventories of bryophytes in mountainous areas of Minas Gerais are still sparse considering the extensive area of the Cerrado domain in the state and in comparison to other terrestrial plant groups. Therefore, Serra da Canastra is an interesting area with great potential for the study of bryophytes (Costa et al. 2011Costa DP, Pôrto KC, Luizi-Ponzo AP, et al. 2011. Synopsis of the Brazilian moss flora: checklist, distribution and conservation. Nova Hedwigia 93: 277-334). Furthermore, there are few compilations of the occurrence of Brazilian bryophytes among phytogeographic domains that also include their distribution within Brazil and worldwide.

To date there has been no study of the bryophytes in this significant biological reserve of Minas Gerais. Therefore, this study aimed to carry out a floristic survey of the bryophytes in Serra da Canastra National Park by analyzing bryophyte species richness, occurrence among phytogeographic domains and Brazilian and worldwide geographical distributions.

Materials and methods

Study area

The Serra da Canastra National Park is situated on a mountain range located in southwestern Minas Gerais and encompasses parts of the municipalities of Sacramento, São Roque de Minas and Delfinópolis (20°00'-20°30'S and 46°15'-47°00'W). It contains 71,525 ha with average altitude ranging from approximately 800 to 1,200 m, with the highest point of the park being Serra Brava at 1,496 m. The average temperature is around 17°C in the winter and 23°C in the summer, but varies between higher and lower regions. Average annual rainfall ranges from approximately 1,300 to 1,700 mm, with most rainfall being concentrated from December to February (IBDF 1981IBDF - Instituto Brasileiro de Desenvolvimento Florestal . 1981. Plano de manejo. Parque Nacional da Serra da Canastra. Brasília, IBDF.).

There are several phytophysiognomies in Serra da Canastra National Park, which provide many conditions for the development of bryophytes, such as rocky fields (campos rupestres), rocky cerrado (cerrado rupestre), grasslands , "dirty fields" (campos sujos), cerrado sensu stricto, gallery forests (matas de galerias), slope forests (matas de encosta), moist fields (campos úmidos), and waterfall areas (Couto Júnior et al. 2010Couto Júnior AFC, Souza VV, Junior OAC, et al. 2010. Integração de parâmetros morfométricos e imagem aster para a delimitação das fitofisionomias da Serra da Canastra, Parque Nacional da Serra da Canastra, MG. Revista Brasileira de Geomorfologia 11: 57- 68. ).

Data collection and identification

Twenty samples from the study area were found in SP herbaria (Holmgren et al. 1981Holmgren PK, Keuken W, Schofield EK. 1981. Index Herbariorum. Part I. The herbaria of the world. 7th. edn. Utrecht/Antwerpen, Bohn, Scheltema & Holkema. ) and four one-week long expeditions were made to the park during to make collections in 2012 to 2014.

Collections were made by walking freely through the main phytophysiognomies of the park including waterfalls, rocky fields (campos rupestres), gallery forest (matas de galeria) and slope forest (matas de encosta) (Couto Júnior et al. 2010Couto Júnior AFC, Souza VV, Junior OAC, et al. 2010. Integração de parâmetros morfométricos e imagem aster para a delimitação das fitofisionomias da Serra da Canastra, Parque Nacional da Serra da Canastra, MG. Revista Brasileira de Geomorfologia 11: 57- 68.). Substrates suitable for bryophyte colonization were classified according to Robbins (1952Robbins RG. 1952. Bryophyte Ecology of a Dune Area in New Zealand. Vegetatio, Acta Geobotanica 4: 1-31. ) and Bates (2009Bates JW. 2009. Mineral nutrition and substratum ecology. In: Goffinet B, Shaw AJ. (eds.) Bryophyte Biology. 2nd. edn. New York, Cambdrige University Press. p. 299-356.) with some modifications. The methodology for collection, herborization and preservation of material followed Gradstein et al. (2001)Gradstein SR , Churchill, SP, Salazar-Allen N. 2001. Guide to the Bryophytes of Tropical America. Memoirs of The New York Botanical Garden 86: 1-577., and all samples collected were deposited in SP.

Identification of the species followed Frahm (1991Frahm JP. 1991. Dicranaceae: Campylopodioideae, Paraleucobryoideae. Flora Neotropica Monograph 54: 1-237. ), Sharp et al. (1994Sharp AJ, Crum H, Eckel P. 1994. The Moss Flora of Mexico. Memoirs of The New York Botanical Garden 69: 1-1113.), Yano & Carvalho (1995Yano O, Carvalho AB. 1995. Briófitas da Serra da Piedade, Minas Gerais, Brasil. In: Anais do 9o Congresso da Sociedade Botânica de São Paulo. São Paulo, Universidade Estadual Paulista. p.15-25.), Buck (1998Buck WR. 1998. Pleurocarpous Mosses of the West Indies. Memoirs of The New York Botanical Garden 1: 1-401. ), Vilas Bôas-Bastos & Bastos (1998Vilas Bôas-Bastos SB, Bastos CJP. 1998. Briófitas de uma área de cerrado no município de Alagoinhas, Bahia, Brasil. Tropical Bryology 15: 101-110.), Bastos et al. (2000)Bastos CJP, Yano O, Vilas Bôas-Bastos SB. 2000. Briófitas de Campos rupestres da Chapada Diamantina, Estado da Bahia, Brasil. Revista Brasileira de Botânica 23: 357-368., Gradstein et al. (2001)Gradstein SR , Churchill, SP, Salazar-Allen N. 2001. Guide to the Bryophytes of Tropical America. Memoirs of The New York Botanical Garden 86: 1-577., Castro et al. (2002Castro NMCF, Pôrto KC, Yano O, Castro AAJF. 2002. Levantamento florístico de bryopsida de cerrado e mata ripícola do parque nacional de Sete Cidades, Piauí, Brasil. Acta Botanica Brasilica 16: 61-76.), Gradstein & Costa (2003), Câmara & Costa (2006Câmara PEAS, Costa DP. 2006. Hepáticas e antóceros das matas de galeria da Reserva Ecológica do IBGE, RECOR, Distrito Federal, Brasil. Hoehnea 33: 79 - 87. ), Câmara (2008aCâmara PEAS. 2008a. Musgos pleurocárpicos das matas de galeria da Reserva Ecológica do IBGE, RECOR, Distrito Federal, Brasil. Acta Botanica Brasilica 22: 573-581. ; bCâmara PEAS. 2008b. Musgos acrocárpicos das Matas de Galeria da Reserva Ecológica do IBGE, RECOR, Distrito Federal, Brasil. Acta Botanica Brasilica 22: 1027-1035. ), Ballejos & Bastos (2009Ballejos J, Bastos CJP. 2009. Musgos Pleurocárpicos do Parque Estadual das Sete Passagens, Miguel Calmon, Bahia, Brasil. Hoehnea 36: 479-495. ), Yano & Peralta (2009)Yano O, Peralta DF. 2009. Flora de Grão-Mogol, Minas Gerais. Briófitas (Bryophyta e Marchantiophyta). Boletim de Botânica da Universidade de São Paulo 27: 1-26., Valente et al. (2011Valente EB, Pôrto KC, Bastos CJP. 2011. Checklist of bryophytes of Chapada Diamantina, Bahia, Brazil. Boletim do Instituto de Botânica 21: 111-124.), Yano & Peralta (2011a; b), Bordin & Yano (2013Bordin J, Yano O. 2013. Fissidentaceae (Bryophyta) do Brasil. Boletim do Instituto de Botânica 22: 1-72. ), Luizi-Ponzo et al. (2013Luizi-Ponzo AP, Siviero TS, Amorim ET, et al 2013. Briófitas do Parque Estadual do Ibitipoca no Herbário Prof. Leopoldo Kriegeer. In: Forzza RC, Neto LM, Salimena FRG, Zappi D. (eds.) Flora do Parque Estadual do Ibitipoca e seu entorno. Vol. 4. 1st. edn. Juiz de Fora, Editora UFJF. p. 95-122.) and Valente et al. (2013)Valente EB, Pôrto KC, Bastos CJP. 2013. Species Richness and Distribution of bryophythes within different phytophysiognomies in the Chapada Diamantina region of Brazil. Acta Botanica Brasilica 27: 294-310..

The classifications followed herein are those of Renzaglia et al. (2009Renzaglia KS, Villarreal JC, Duff RJ. 2009. New insights into morphology, anatomy and systematics of hornworts. In: Goffinet B, Shaw AJ. (eds.) Bryophyte Biology . 2nd. edn. New York, Cambdrige University Press . p. 139-171.) for Anthocerotophyta, Crandall-Stotler et al. (2009Crandall-Stotler B, Stotler RE, Long DG. 2009. Morphology and classification of the Marchantiophyta. In: Goffinet B, Shaw AJ. (eds.) Bryophyte Biology . 2nd. edn. New York, Cambdrige University Press . p. 1-54.) for Marchantiophyta and Goffinet et al. (2009Goffinet B, Buck WR, Shaw AJ. 2009. Morphology, anatomy and classification of the Bryophyta. In: Goffinet B, Shaw AJ. (eds.) Bryophyte Biology . 2nd. edn. New York, Cambdrige University Press . p. 56-138.) for Bryophyta.

The list of species is organized in alphabetical order by family, then genus and then species, and provides the Brazilian geographical distribution as proposed by Valente & Pôrto (2006Valente EB, Pôrto KC. 2006. Hepáticas (Marchantiophyta) de um fragmento de Mata Atlântica na Serra da Jibóia, município de Santa Terezinha, BA, Brasil. Acta Botanica Brasilica 20: 433-441.), who considered as "rare" those bryophythes occurring in one to four Brazilian states; as "moderate" those occurring in five to nine states; and as "wide" those occurring in ten or more Brazilian states. The distribution and biogegraphical domains followed Gradstein & Costa (2003Gradstein SR, Costa DP. 2003. The Hepaticae and Anthocerotae of Brazil. Memoirs of The New York Botanical Garden 87: 1-318. ), Yano (2008Yano O. 2008. Catálogo de antóceros e hepáticas brasileiros: literatura original, basiônimo, localidade-tipo e distribuição geográfica. Boletim do Instituto de Botânica 19: 1-110. ), Forzza et al. (2010Forzza RC, Leitman PM, Costa AF, et al. 2010. Introdução. In: Forzza RC, Baumgratz JFA, Bicudo CEM, et al. (eds.) Lista de Espécies da Flora do Brasil. Jardim Botânico do Rio de Janeiro . Vol. 1. Rio de Janeiro, Jardim Botânico do Rio de Janeiro. ), Costa et al. (2011)Costa DP, Pôrto KC, Luizi-Ponzo AP, et al. 2011. Synopsis of the Brazilian moss flora: checklist, distribution and conservation. Nova Hedwigia 93: 277-334 and Costa & Peralta (2015)Costa DP, Peralta DF. 2015. Briófitas In: Lista de Espécies da Flora do Brasil. Jardim Botânico do Rio de Janeiro. http://floradobrasil.jbrj.gov.br/jabot/floradobrasil/FB128472. 19 Aug. 2015.
http://floradobrasil.jbrj.gov.br/jabot/f...
.

Results and discussion

Species richness

One thousand one hundred and seventy-seven (1,177) samples were analyzed, with 722 samples of mosses (Bryophyta), 454 of liverworts (Marchantiophyta) and only one of a hornwort (Anthocerotophyta). Two hundred and eight-nine species of bryophytes were found, representing 128 genera and 57 families, accounting for approximately 38% of the species estimated for the state and 19% for the country (Tab. 1) (Costa & Peralta 2015Costa DP, Bastos CJP, Schäfer-Verwimp A. 2015. Lejeuneaceae In: Lista de Espécies da Flora do Brasil. Jardim Botânico do Rio de Janeiro. http://www.floradobrasil.jbrj.gov.br/jabot/floradobrasil/FB97355. 19 Aug. 2015.
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).

The family Lejeuneaceae was the most represented with 53 species (18%). The number of moss species was greater (174 species) than that of liverworts (114 species), while only one species of hornwort was found (Tab. 1). Among mosses (Bryophyta) the most common families were Sphagnaceae, with 26 species, followed by Leucobryaceae (21 species), Bryaceae (13 species) and Sematophyllaceae (12 species). Among liverworts (Marchantiophyta), the most common family was Lejeuneaceae, with 53 species, followed by Lepidoziaceae (10 species) and Frullaniaceae (nine species). Fifty-six species (19%) were considered new ocurrences in the state, while 14% of bryophytes species (41 species) were considered endemic to Brazil.

Table 1
List of species found in Serra da Canastra National Park. Phytogeographic domains (Phyt. dom.): Amazon Rainforest = AM, Atlantic Forest = AT, Cerrado = CE, Caatinga = CA, Pampa = PA, Pantanal = PL; Geographical distribution of Brazil (Brazil distr.); Geographical distribution worldwide (World distr.); * = New record for Minas Gerais State.

Other studies in high altitude areas of Minas Gerais State, also found greater species richness of mosses than liverworts (Yano & Carvalho 1995Yano O, Carvalho AB. 1995. Briófitas da Serra da Piedade, Minas Gerais, Brasil. In: Anais do 9o Congresso da Sociedade Botânica de São Paulo. São Paulo, Universidade Estadual Paulista. p.15-25.; Yano & Peralta 2009Yano O, Peralta DF. 2009. Flora de Grão-Mogol, Minas Gerais. Briófitas (Bryophyta e Marchantiophyta). Boletim de Botânica da Universidade de São Paulo 27: 1-26.; Yano & Peralta 2011aYano O, Peralta DF. 2011a. Bryophytes from Serra de São José, Tiradentes, Minas Gerais, Brasil. Boletim de Botânica da Universidade de São Paulo 21: 141-172.; bYano O, Peralta DF. 2011b. Flora da Serra do Cipó, Minas Gerais: Briófitas (Anthocerotophyta, Bryophyta e Marchantiophyta). Boletim de Botânica da Universidade de São Paulo 29: 135-211.; Luizi-Ponzo et al. 2013Luizi-Ponzo AP, Siviero TS, Amorim ET, et al 2013. Briófitas do Parque Estadual do Ibitipoca no Herbário Prof. Leopoldo Kriegeer. In: Forzza RC, Neto LM, Salimena FRG, Zappi D. (eds.) Flora do Parque Estadual do Ibitipoca e seu entorno. Vol. 4. 1st. edn. Juiz de Fora, Editora UFJF. p. 95-122.). This is probably due to mosses having more morphologically complex gametophytes and sporophytes, which increases the possibilities for occupation of varied environments. Furthermore, mosses are more resistant to dessication than most of liverworts (Goffinet et al. 2009Goffinet B, Buck WR, Shaw AJ. 2009. Morphology, anatomy and classification of the Bryophyta. In: Goffinet B, Shaw AJ. (eds.) Bryophyte Biology . 2nd. edn. New York, Cambdrige University Press . p. 56-138.), thereby they can occur and predominate in areas where the availability of water is more restricted, and conditons of moisture and rainfall are relatively low, unlike the Atlantic Forest where the species richness of mosses is lower than that of liverworts (Gradstein et al. 2001Gradstein SR , Churchill, SP, Salazar-Allen N. 2001. Guide to the Bryophytes of Tropical America. Memoirs of The New York Botanical Garden 86: 1-577.).

Sphagnaceae was the most species rich moss family and, in including a number of Brazilian endemics and new occurrences, of significant importance to Serra da Canastra (Tab. 1). Although this family is often associated with watercourses and water-soaked soils (Schofield 1985Schofield WB. 1985. Introduction to Bryology. New York, Macmillan Publisching Company. ), it can also be found on rocky and extremely dry surfaces, with soil and rocks as their main substrates.

Leucobryaceae (21 species) and Bryaceae (13 species) were the bryophyte families with the greatest richness in high altitudes areas in Minas Gerais (Yano & Carvalho 1995Yano O, Carvalho AB. 1995. Briófitas da Serra da Piedade, Minas Gerais, Brasil. In: Anais do 9o Congresso da Sociedade Botânica de São Paulo. São Paulo, Universidade Estadual Paulista. p.15-25.; Yano & Peralta 2011bYano O, Peralta DF. 2011b. Flora da Serra do Cipó, Minas Gerais: Briófitas (Anthocerotophyta, Bryophyta e Marchantiophyta). Boletim de Botânica da Universidade de São Paulo 29: 135-211.). This is supported by the fact that representatives of these families are constantly found in mountain regions (Gradstein et al. 2001Gradstein SR , Churchill, SP, Salazar-Allen N. 2001. Guide to the Bryophytes of Tropical America. Memoirs of The New York Botanical Garden 86: 1-577.). These families have in common an acrocarpous growth habit of the gametophyte, and are found in more exposed areas, such as the vast rocky fields of the Cerrado (Bastos & Vilas Bôas-Bastos 2008Bastos CJP, Vilas Bôas-Bastos SB. 2008. Musgos Acrocárpicos e Cladocárpicos (Bryophyta) da Reserva Ecológica da Michelin, Igrapiúna, Bahia, Brasil. Stientibus Série Ciências Biológicas 8: 275-279. ). Sematophyllaceae and Hypnaceae were the most represented pleurocarpous moss families and are widely distributed worldwide, mainly in the tropics, and are very diverse morphologically (Buck 1998Buck WR. 1998. Pleurocarpous Mosses of the West Indies. Memoirs of The New York Botanical Garden 1: 1-401. ; Ramsay et al. 2002Ramsay HP, Schofield WB, Tan BC. 2002. The family Sematophyllaceae (Bryopsida) in Australia. Part 1: Introduction, family data, key to genera and the genera Wijkia, Acanthorrynchium, Trismegistia and Sematophyllum. The Journal of Hattori Botanical Laboratory 92: 1-50. ; Ireland & Buck 2009Ireland RR, Buck WR. 2009. Some Latin American Genera of Hypnaceae. Smithsonian Contributions to Botany 93: p. 1-97.).

The family of liverworts with the greatest species richness was Lejeuneaceae, which has 56 genera and 285 species in Brazil (Costa et al. 2015Costa DP, Bastos CJP, Schäfer-Verwimp A. 2015. Lejeuneaceae In: Lista de Espécies da Flora do Brasil. Jardim Botânico do Rio de Janeiro. http://www.floradobrasil.jbrj.gov.br/jabot/floradobrasil/FB97355. 19 Aug. 2015.
http://www.floradobrasil.jbrj.gov.br/jab...
). In the present study the family accounted for 47% of the liverworts found, and has frequently been among the most abundant families of liverworts in other studies on mountainous areas of Minas Gerais (Yano & Carvalho 1995Yano O, Carvalho AB. 1995. Briófitas da Serra da Piedade, Minas Gerais, Brasil. In: Anais do 9o Congresso da Sociedade Botânica de São Paulo. São Paulo, Universidade Estadual Paulista. p.15-25.; Yano & Peralta 2009Yano O, Peralta DF. 2009. Flora de Grão-Mogol, Minas Gerais. Briófitas (Bryophyta e Marchantiophyta). Boletim de Botânica da Universidade de São Paulo 27: 1-26.; Yano & Peralta 2011aYano O, Peralta DF. 2011a. Bryophytes from Serra de São José, Tiradentes, Minas Gerais, Brasil. Boletim de Botânica da Universidade de São Paulo 21: 141-172.; bYano O, Peralta DF. 2011b. Flora da Serra do Cipó, Minas Gerais: Briófitas (Anthocerotophyta, Bryophyta e Marchantiophyta). Boletim de Botânica da Universidade de São Paulo 29: 135-211.; Luizi-Ponzo et al. 2013Luizi-Ponzo AP, Siviero TS, Amorim ET, et al 2013. Briófitas do Parque Estadual do Ibitipoca no Herbário Prof. Leopoldo Kriegeer. In: Forzza RC, Neto LM, Salimena FRG, Zappi D. (eds.) Flora do Parque Estadual do Ibitipoca e seu entorno. Vol. 4. 1st. edn. Juiz de Fora, Editora UFJF. p. 95-122.). This family is known to have originated relatively recently during bryophyte evolution, and includes many independent diversifications resulting in great morphological variation, several forms of substrate occupation and different taxonomic conceptulizations which have resulted in this great diversity presented by the family (Groth-Malonek et al. 2004Groth-Malonek M, Heinrichs J, Schneider H, Gradstein SR. 2004. Philogenetic relationships in the Lejeuneaceae (Hepaticae) inferred using ITS sequences of nuclear ribosomal DNA. Organisms, Diversity and Evolution 4: 51-57. ).

Phaeoceros laevis was the only species of hornwort (Anthocerotophyta) found in Serra da Canastra National Park, where it was collected growing on rocks among rocky outcrops near streams of the lower portion of the Casca D'anta waterfall. Only one of the previous studies in Minas Gerais also found this species (Yano & Peralta 2011bYano O, Peralta DF. 2011b. Flora da Serra do Cipó, Minas Gerais: Briófitas (Anthocerotophyta, Bryophyta e Marchantiophyta). Boletim de Botânica da Universidade de São Paulo 29: 135-211.). In Serra do Cipó, this species was found growing on soil and was associated with the thallose liverwort Fossombronia porphyrorhyza.

Substrate colonization

Regarding substrate colonization, a predominance of rupicolous bryophytes was observed in Serra da Canastra National Park. Among the 1,177 bryophyte samples collected, 406 were on rocky substrates, 356 were corticolous, 349 terrestrial, 27 epiphyllous and 18 epixylic (Fig. 1). Mosses that were found growing submerged or under the water were considered aquatic and, with other different substrates types (termite mound, litter), represented 21 samples (Fig. 1).

In other studies conducted in Minas Gerais, species of bryophytes showed a preference for corticolous substrates (Yano & Peralta 2011bYano O, Peralta DF. 2011b. Flora da Serra do Cipó, Minas Gerais: Briófitas (Anthocerotophyta, Bryophyta e Marchantiophyta). Boletim de Botânica da Universidade de São Paulo 29: 135-211.; Luizi-Ponzo et al. 2013Luizi-Ponzo AP, Siviero TS, Amorim ET, et al 2013. Briófitas do Parque Estadual do Ibitipoca no Herbário Prof. Leopoldo Kriegeer. In: Forzza RC, Neto LM, Salimena FRG, Zappi D. (eds.) Flora do Parque Estadual do Ibitipoca e seu entorno. Vol. 4. 1st. edn. Juiz de Fora, Editora UFJF. p. 95-122.), such as growing on the trunks of living trees. In their study focusing on gallery forests of Serra do Cipó, Souza & Câmara (2015Souza RV, Câmara PEAS. 2015. Survey of the bryophytes of a gallery forest in the National Park of Serra do Cipó, Minas Gerais, Brazil. Acta Botanica Brasilica 29: 24-29.) also observed a preference for corticicolous substrates and explanained this finding on the abundance of this substrate in the area, but concluded that ecological studies are needed to verify this hypothesis. Following this reasoning, the large amount of rupicolous species in Serra da Canastra National Park is likely due to the presence of many rocks, especially in areas near waterfalls and in rocky fields, since the methodology of collection was the same for all vegetation types. However, as with the gallery forest of Serra do Cipó, ecological study is necessary to conclude this hypothesis.

Our study collected 27 samples (Fig. 1) of epiphyllous bryophytes, which are those species that grow on living leaves. The family Lejeuneaceae was the only family found growing on this type of substrate in Serra da Canastra National Park. According to Gradstein et al. (2001)Gradstein SR , Churchill, SP, Salazar-Allen N. 2001. Guide to the Bryophytes of Tropical America. Memoirs of The New York Botanical Garden 86: 1-577., this family can colonize many types of substrates, but the epiphyllous habit is common in preserved native areas, which highlights the exigency for the protection of Serra da Canastra National Park.

Figure 1
Percentage of samples obtained from each substrate.

Worldwide and Brazilian distribution

Concerning worldwide distribution, a predominance of Neotropical species was observed in Serra da Canastra National Park, with 89 species. This was followed by 61 tropical America species, 41 Brazilian endemic species, 39 cosmopolitan species, 33 pantropical species and 26 with disjunct distributions with the Andes, Africa and other countries of South America (Fig. 2) (Tab. 1).

Figure 2
Percentage of bryophytes species obtained and their worldwide distribution.

Most of these species (132) have a wide geographic distribution in Brazil, accounting for 46% of bryophytes found in the area (Tab. 1). These species are typically generalists and easily occupy a variety of environments and several types of substrates. Examples of such mosses (Bryophyta) include Bryum argenteum, Hyophila involuta, Octoblepharum albidum, Sematophyllum subpinnatum and Syrrhopodon prolifer. Example of such liverworts (Marchantiophyta) include Frullania ericoides and Lejeunea flava (Forzza et al. 2010Forzza RC, Leitman PM, Costa AF, et al. 2010. Introdução. In: Forzza RC, Baumgratz JFA, Bicudo CEM, et al. (eds.) Lista de Espécies da Flora do Brasil. Jardim Botânico do Rio de Janeiro . Vol. 1. Rio de Janeiro, Jardim Botânico do Rio de Janeiro. ). Ninety species (31%) had a moderate distribution and 67 (23%) were considered rare (Tab. 1).

We found 56 new species for the state of Minas Gerais, which represents 19% of the total bryophyte species collected. Among these, 36 species (64%) have a "rare" distribution in Brazil, 13 (23%) a "moderate" distribution and seven (13%) a "wide" distribution (Tab. 1). Campylopus densicoma was first recorded for Minas Gerais and has a "rare" distribution, being previously found only in Rio de Janeiro, and is considered an endangered species (Costa et al. 2013Costa DP, Fernandez EP, Monteiro NP, Messina T. 2013. Dicranaceae. In: Martinelli G, Moraes MA. (eds.) Livro Vermelho da Flora do Brasil. Rio de Janeiro, CNCFLORA. p. 477-478.).

The data obtained through this study show that the Serra da Canastra National Park is an important area for bryophyte diversity and conservation, especially since there remain many species to be discovered.

Phytogeographic domains

Serra da Canastra National Park is located in the Cerrado domain (IBDF 1981IBDF - Instituto Brasileiro de Desenvolvimento Florestal . 1981. Plano de manejo. Parque Nacional da Serra da Canastra. Brasília, IBDF.). However, most of the species (62%) found in the park are considered exclusive to the Atlantic Forest domain, such as Bryum caespiticium, Squamidium brasiliense, Syrrhopodon tortilis and Wijkia flagellifera for Bryophyta (Souza & Câmara 2015Souza RV, Câmara PEAS. 2015. Survey of the bryophytes of a gallery forest in the National Park of Serra do Cipó, Minas Gerais, Brazil. Acta Botanica Brasilica 29: 24-29.) and Bazzania heterostipa, Calypogeia grandistipula, Frullania schaefer-verwimpii and Lejeunea cristulata for Marchantiophyta (Yano 2008Yano O. 2008. Catálogo de antóceros e hepáticas brasileiros: literatura original, basiônimo, localidade-tipo e distribuição geográfica. Boletim do Instituto de Botânica 19: 1-110. ).

Moreover, the present study also found some Brazilian endemic species, which had so far been only recorded for the Amazon Rainforest domain, such as Cololejeunea contractiloba and Lejeunea erostrata of Marchantiophyta, and Sphagnum dimorphophyllum for Bryophyta (Costa et al. 2011Costa DP, Pôrto KC, Luizi-Ponzo AP, et al. 2011. Synopsis of the Brazilian moss flora: checklist, distribution and conservation. Nova Hedwigia 93: 277-334). Perhaps these results are due to gaps in collections of material from areas of the Brazilian Highlands, such as Serra da Canastra (Gradstein et al. 2001Gradstein SR , Churchill, SP, Salazar-Allen N. 2001. Guide to the Bryophytes of Tropical America. Memoirs of The New York Botanical Garden 86: 1-577.; Costa et al. 2011Costa DP, Pôrto KC, Luizi-Ponzo AP, et al. 2011. Synopsis of the Brazilian moss flora: checklist, distribution and conservation. Nova Hedwigia 93: 277-334), which is reflected in the great number of new occurrences observed in this study.

Some typical Cerrado species were found, such as Riccardia regnelli, Mesonodon regnellianus, Campylopus dichrostis, Leucobryum clavatum and Jonesiobryum cerradense, as well as some species exclusive to the Cerrado, for example, Itatiella denudata, Sphagnum delamboyense and Sphagnum garysmithii (Costa et al. 2011Costa DP, Pôrto KC, Luizi-Ponzo AP, et al. 2011. Synopsis of the Brazilian moss flora: checklist, distribution and conservation. Nova Hedwigia 93: 277-334).

The two species endemic to Serra da Canastra National Park, Itatiella denudata and Sphagnum geraisense, were recollected during this study reinforcing the importance of maintaining Serra da Canastra National Park for the conservation of endemic species (Romero & Nakajima 1999Romero R, Nakajima JN. 1999. Espécies endêmicas doParque Nacional da Serra da Canastra , Minas Gerais. Revista Brasileira de Botânica 22: 259-265) as well as the great diversity of other bryophytes found in this study.

This study has provided new information on bryophyte richness, occurrence, occupied phytophysiognomies and worldwide and Brazilian distributions, particularly those occurring in mountain areas of the Brazilian Highlands, where more floristic studies on bryophytes are needed. The large number of species found in the park (289 spp.), accounting for approximately 38% of the species known in Minas Gerais and 19% in Brazil, the new records for this state (56 spp.) and the number of Brazilian endemic species (41 spp.), stress the importance of performing floristic inventories in Brazil and maintaining Serra da Canastra National Park for conservation as a biological reserve.

Acknowledgments

We acknowledge the Botanical Institute of São Paulo (Instituto Botânico de São Paulo) and the Bryology Section (Seção de Briologia) for the support of all the necessary equipment throughout the study. We also acknowledge CAPES for financial support and ICMBIO for the license n° 39601-2 granted collecting in the area.

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Publication Dates

  • Publication in this collection
    19 Apr 2016
  • Date of issue
    Apr-Jun 2016

History

  • Received
    09 Sept 2015
  • Accepted
    23 Mar 2016
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