An illustrated guide of ferns and lycophytes from Carambeí, PR, Brazil

Cássio Michelon Frederico Fregolente Faracco Mazziero Bianca Kalinowski Canestraro Mathias Erich Engels About the authors

Abstract

We present a list of species of ferns and lycophytes from Carambeí, a municipality located in Paraná state, southern Brazil. This area, locally known as “Campos Gerais”, presents an intricate mosaic of forests and savannah with several canyons and humid valleys that harbors a rich and unique vegetation in Southern Brazil. In total, we found 129 species (119 ferns and 10 lycophytes), distributed in 18 families and 59 genera, and 20% (26 spp.) of which are endemic to Brazil. The most expressive families were Polypodiaceae (19 spp.), Pteridaceae (18 spp.) and Thelypteridaceae (15 spp.). The most representative genera were Asplenium (8 spp. - 6.2%) and Amauropelta (8 spp - 6.2%). Terrestrial herbs, epiphytes and lithophytes are the most common guild of life forms, successively. The richest environments were woody forest (61% of the species) and herbaceous-shrubby (12%). Arachniodes denticulata, Phlegmariurus flexibilis and P. Heterocarpon are the second record in the “Campos Gerais” region. We also provide photographic plates with diagnostic characters for most of the species.

Key words:
Campos Gerais; diversity; floristic; pteridophytes; species richness

Resumo

É apresentada uma listagem de espécies de samambaias e licófitas para o município de Carambeí, localizado no estado do Paraná, Sul do Brasil. Essa área, localmente conhecida como Campos Gerais, apresenta um complexo mosaico de florestas e Cerrado com diversos canyons e vales úmidos e comporta uma vegetação rica e singular no Sul do Brasil. No total, foram registradas 129 espécies (119 samambaias e 10 licófitas), distribuídas em 18 famílias e 59 gêneros, sendo 20% (26 spp.) endêmicas do Brasil. As famílias mais expressivas são Polypodiaceae (19 spp.), Pteridaceae (18 spp.) e Thelypteridaceae (15 spp.). Os gêneros mais representativos foram Asplenium (8 spp. - 6,2%) e Amauropelta (8 spp. - 6,2%). As guildas de formas de vida mais comuns são ervas terrícolas, epífitas e rupícolas, sucessivamente. Os ambientes mais ricos foram os florestais (61% das espécies) e os herbáceo-arbustivos (12%). Arachniodes denticulata, Phlegmariurus flexibilis e P. heterocarpon foram registradas pela segunda vez para a região dos Campos Gerais. São fornecidas pranchas fotográficas com caracteres diagnósticos para a maioria das espécies.

Palavras-chave:
Campos Gerais; diversidade; florística; pteridófitas; riqueza de espécies

Introduction

Ferns and lycophytes are an important component of the vascular flora in tropical forests, representing 10% to 14% of the diversity in these environments, often being the dominant groups in the understory (Gentry 1990Gentry AH (1990) Floristic similarities and differences between Southern Central America and upper and Central Amazonia. In: Gentry AH (ed.) Four neotropical rain forests. Yale University Press, New Haven. Pp. 141-160.; Costa 2004Costa FRC (2004) Structure and composition of the ground-herb community in a terra-firme Central Amazonian forest. Acta Amazonica 34: 53-59.; Moran 2008Moran RC (2008) Diversity, biogeography and floristics. In: Ranker TA & Haufler CH (eds.) Biology and evolution of ferns and lycophytes. Cambridge University Press, Cambridge. Pp. 367-394.). In Brazil, these groups are represented by 1,318 species, 503 (38%) of which are endemic (Prado et al. 2015Prado J, Sylvestre LS, Labiak PH, Windisch PG, Salino A, Barros ICL, Hirai RY, Almeida TE, Santiago ACP, Kieling-Rubio MA, Pereira AFN, Øllgaard B, Ramos CG, Mickel JT, Dittrich VAO, Mynssen CM, Schwartsburd PB, Condack JPS, Pereira JBS & Matos FB (2015) Diversity of ferns and lycophytes in Brazil. Rodriguésia 66: 1073-1083.). The Atlantic Forest is the most diverse domain in Brazil harboring 883 species and the Ombrophilous Forest is the type of vegetation with greatest diversity with 90% of the species in this domain (Prado et al. 2015Prado J, Sylvestre LS, Labiak PH, Windisch PG, Salino A, Barros ICL, Hirai RY, Almeida TE, Santiago ACP, Kieling-Rubio MA, Pereira AFN, Øllgaard B, Ramos CG, Mickel JT, Dittrich VAO, Mynssen CM, Schwartsburd PB, Condack JPS, Pereira JBS & Matos FB (2015) Diversity of ferns and lycophytes in Brazil. Rodriguésia 66: 1073-1083.). This diversity is reflected in several studies carried out in these areas, such as, Mynssen & Windisch (2004)Mynssen CM & Windisch PG (2004) Pteridófitas da Reserva Rio das Pedras, Mangaratiba, RJ, Brasil. Rodriguesia 55: 125-156. for Rio de Janeiro state, Salino & Almeida (2008)Salino A & Almeida TE (2008) Pteridófitas do Parque Estadual do Jacupiranga, SP, Brasil. Acta Botanica Brasilica 22: 983-991. and Mazziero et al. (2015)Mazziero FFF, Labiak PH & Paciencia M (2015) Checklist of ferns and lycophytes from the Parque Estadual Turístico do Alto Ribeira, Iporanga, São Paulo, Brazil. Check List 11: 1791. for São Paulo state and Matos et al. (2010)Matos FB, Amorim AM & Labiak PH (2010) The ferns and lycophytes of a montane tropical forest in southern Bahia, Brazil. Journal of the Botanical Research Institute of Texas 4: 333-346. for Bahia state.

In the state of Paraná, the areas of Atlantic Rain Forests in the coastal region are responsible for a high richness of ferns and lycophytes (Prado et al. 2015Prado J, Sylvestre LS, Labiak PH, Windisch PG, Salino A, Barros ICL, Hirai RY, Almeida TE, Santiago ACP, Kieling-Rubio MA, Pereira AFN, Øllgaard B, Ramos CG, Mickel JT, Dittrich VAO, Mynssen CM, Schwartsburd PB, Condack JPS, Pereira JBS & Matos FB (2015) Diversity of ferns and lycophytes in Brazil. Rodriguésia 66: 1073-1083.), which is strongly related with the great diversity of available environments in mountainous areas (Moran 1995Moran RC (1995a) The importance of mountains to pteridophytes, with emphasis on neotropical rain forests. In: Churchill SP, Balslev H, Forero E & Luteyn JL (eds.) Biodiversity and conservation on neotropical montane forests. New York Botanical Garden, New York. Pp. 359-563.). In contrast, in the inland of Paraná, the relief of the “Campos Gerais” region is not mountainous, but it presents a unique arrange of geomorphologic characteristics (Maack 1981Maack R (1981) Geografia física do estado do Paraná. José Olympio, Rio de Janeiro. 450p.). Recent studies carried out in these areas (Schwartsburd and Labiak 2007Schwartsburd PB, Labiak PH & Salino A (2007) A new species of Ctenitis (Dryopteridaceae) from southern Brazil. Brittonia 59: 29-32.; Michelon & Labiak 2013Michelon C & Labiak PH (2013) Samambaias e licófitas do Parque Estadual do Guartelá, PR, Brasil. Hoehnea 40: 191-204.) have shown the importance of inland regions to the diversity of ferns and lycophytes as well. Nevertheless, few studies have been performed in the inland portion of Paraná, for example Sakagami (2006)Sakagami CR (2006) Pteridófitas do Parque Ecológico da Klabin, Telêmaco Borba, Paraná, Brasil. Dissertação de Mestrado, Universidade Federal do Paraná, Curitiba. 212p., Schwartsburd & Labiak (2007)Schwartsburd PB & Labiak PH (2007) Pteridófitas do Parque Estadual de Vila Velha, Ponta Grossa, Paraná, Brasil. Hoehnea 34: 159-209., Michelon & Labiak (2013)Michelon C & Labiak PH (2013) Samambaias e licófitas do Parque Estadual do Guartelá, PR, Brasil. Hoehnea 40: 191-204., Lautert et al. (2015)Lautert M, Temponi LG, Viveros RS & Salino A (2015) Lycophytes and ferns composition of Atlantic Forest conservation units in western Paraná with comparisons with other areas in southern Brazil. Acta Botanica Brasilica 29: 499-508. and, new species of ferns from this region have been recently described (Schwartsburd et al. 2007Schwartsburd PB & Labiak PH (2007) Pteridófitas do Parque Estadual de Vila Velha, Ponta Grossa, Paraná, Brasil. Hoehnea 34: 159-209.; Schwartsburd & Labiak 2008Schwartsburd PB & Labiak PH (2008) Eriosorus areniticola (Pteridaceae), a new species from Brazil. American Fern Journal 98: 160-163.; Christenhusz et al. 2009Christenhusz MJM, Schwartsburd PB & Labiak PH (2009) Hymenophyllum filmenofilicum (Hymenophyllaceae: Pteridophyta): a new epipetric filmy fern from Paraná, southern Brazil. Kew Bulletin 64: 175-178.). Therefore, floristics studies and plant collections are still crucial considering the distinct mosaic and the high diversity of plants in these areas. It is important to mention that most of these studied areas are conservation units with minor human impacts. Recent data show that there is left no more than 12% of Atlantic Forest remnants in Paraná state, and with very few inland forested areas (Fundação SOS Mata Atlântica & INPE 2016Fundação SOS Mata Atlântica & INPE - Instituto Nacional de Pesquisas Espaciais (2001) Atlas dos remanescentes florestais da Mata Atlântica e ecossistemas associados no período de 1995-2000. Fundação SOS Mata Atlântica e INPE, São Paulo. 61p.). The inland portions of the state suffered a severe deforestation due to agriculture and livestock activities.

This study aims to contribute to the knowledge of the diversity of these groups in the “Campos Gerais” region, providing a list of species, as well as data about geographic distribution, phytophysiognomy and guild of life forms. We also provide photographic records with diagnosis characters for most of the species.

Materials and Methods

Study area

Carambeí is located in Paraná state, southern Brazil (24°57’04”S, 50°06’37”W). Its territory is about 650 km2 occupying the “Campos Gerais” region in the Paraná’s First and Second Plateaus (Maack 1981Maack R (1981) Geografia física do estado do Paraná. José Olympio, Rio de Janeiro. 450p.; Cordeiro Santos et al. 2009Cordeiro Santos LJ, Oka-Fiori C, Canali NE, Fiori AP, Da Silveira CT & Franca Da Silva JM (2009) Morphostructural mapping of Parana state, Brazil. Journal of Maps 5: 170-178.; Labiak 2014aLabiak PH (2014a) Aspectos fitogeográficos do Paraná. In: Kaehler M, Goldenberg R, Labiak PH, Ribas OS, Vieira AOS & Hatschbach GG (eds.) Plantas vasculares do Paraná. Ed. UFPR, Curitiba. Pp. 7-22.). The altitude ranges from 780 m a.s.l., in the proximities of the Tibagi river, to 1,100 m a.s.l., at the top of the Devonian Steep.

In the Paraná’s First Plateau predominates the Araucaria forest and small patches of intensively disturbed grasslands. In the Second Plateau, associated with capon forests, the grasslands dominate the landscape. Also, Cerrado relicts (Brazilian savannah) can be found in the São João river canyon (Maack 1981Maack R (1981) Geografia física do estado do Paraná. José Olympio, Rio de Janeiro. 450p.).

Field work

Collections were performed during March 2013. All possible habitats were explored. In this expedition, the specimens were collected and herborized following the usual methods for the studied groups (Fidalgo & Bononi 1984Fidalgo O & Bononi VLR (1984) Técnicas de coleta, preservação e herborização de material botânico. Manual do Instituto de Botânica nº 4, São Paulo. 62p.) and were deposited in the UPCB herbarium.

Taxonomic treatment

Classification system followed PPG I (2016)PPG I (2016) A community-derived classification for extant lycophytes and ferns. Journal of Sytematics Evolution 54: 563-603.. Species’ authors and abbreviations followed IPNI (2017). Identifications were made by comparison with collections at UPCB, specialists and specific literature. The guilds of life forms were based on Paciencia (2008)Paciencia MLB (2008) Diversidade de pteridófitas em gradientes de altitude na Mata Atlântica do estado do Paraná, Brasil. Tese de Doutorado. Universidade de São Paulo, São Paulo. 229p..

Geographical distribution

Geographical distribution data were obtained from Prado et al. (2015)Prado J, Sylvestre LS, Labiak PH, Windisch PG, Salino A, Barros ICL, Hirai RY, Almeida TE, Santiago ACP, Kieling-Rubio MA, Pereira AFN, Øllgaard B, Ramos CG, Mickel JT, Dittrich VAO, Mynssen CM, Schwartsburd PB, Condack JPS, Pereira JBS & Matos FB (2015) Diversity of ferns and lycophytes in Brazil. Rodriguésia 66: 1073-1083., floras and review papers. The geographic patterns adopted were: Pantropic, Neotropic, South America, Brazil and Southern/Southeastern Brazil.

Vegetation

Regarding the environments in which the species were found, we considered three situations:

Aquatic vegetation:

Alagados reservoir (Fig. 1a): floating vegetation in the Pitanguí hydroelectric power plant reservoir.

Figure 1
Aquatic and woody vegetation – a. Alagados’ reservoir; b. secondary forest border; c. capon forest understory; d,e. gallery forest understory; f. riparian forest understory.

Woody vegetation: arboreal vegetation, mid to large size, primitive (primary forests) or secondary (disturbed forests and capoeirões), divided into:

Secondary forests (Fig. 1b,c): secondary vegetation from initial to medium regeneration stage.

Gallery forests (Fig. 1d,e): forests growing on geological fissures, composed by primary vegetation, usually with rocky streams inside.

Riparian forests (Figs. 1f; 2a): forests that grow along the major rivers in the area (Jotuva, Pitanguí, São João, Tamanduá and Tibagi rivers).

Figure 2
Figure 2 – Woody and herbaceous-shrubby vegetation – a. riparian forest; b. riparian slabstone environment; c. dry grasslands; d. litholic grasslands; e. human disturbed environment; f. waterfall amidst grasslands.

Riparian slabstone environments (Fig. 2b): midsize vegetation growing alongside stony rivers amid grasslands areas, usually in the expansions of the rock-forming slabstone.

Swamp areas: midsize woody vegetation in areas of poorly drained soil, usually amid grasslands, however, without undergoing periodic flooding.

Herbaceous and shrubby vegetation: small vegetation, herbaceous and/or shrubby, primitive (grasslands) or secondary (capoeirinhas), divided into:

Dry grasslands (Fig. 2c): including areas of campo limpo, where the herbaceous vegetation predominates amongst rare woody species, and campo sujo, where the shrubby and woody species are commonly found. Remnants of Cerrado (Brazilian Savannah) are also included in this category.

Litholic grasslands (Fig. 2d): herbaceous and shrubby vegetation occurring in areas of rocky fields with sandstone outcrops.

Human disturbed environment (Fig. 2e): characterized by intensively modified and mostly herbaceous vegetation, such as urbanized areas and roadsides slopes.

Wet grasslands: mainly herbaceous vegetation in areas of poor water drainage (Fig. 2f).

Results

In the present work we found 129 species (119 ferns and 10 lycophytes), distributed in 18 families and 59 genera (Tab. 1 - see supplementary material at <https://doi.org/10.6084/m9.figshare.6142595.v1>). The richest families were Polypodiaceae (19 species - 14.7%), Pteridaceae (18 spp. - 13.9%), Thelypteridaceae (15 spp. - 11.6%), Dryopteridaceae (13 spp. - 10%) and Hymenophyllaceae (12 spp. - 9.3%), which together comprise almost 60% of the total richness. The most expressive genera were Asplenium (8 species - 6.2%), Amauropelta (8 species - 6.2%) and, Doryopteris, Hymenophyllum and Pecluma (6 species - 4.6% each) (Tab. 1).

Table 1
List of ferns and lycophytes recorded in Carambeí municipality, southern Brazil. The life forms are represented by: T = terrestrial herb, E = epiphyte, L = lithophyte, C = climbing, A = terrestrial arborescent, H = hemiepiphyte, Q = aquatic; vegetation: SF = secondary forest, DE = human disturbed environment, LG = litholic grassland, GF = gallery forest, RF = riparian forest, SW = swamp areas, RS = riparian slabstone environment, DG = dry grassland, WG = wet grassland, RE = Alagados reservoir; geographical distribution: NEO = Neotropical, SAM = South American, PAN = Pantropical, BRA = Brazilian, SSE = southern/southeastern Brazilian. Voucher (voucher material at UPCB herbarium: # - Michelon, C. collector number, * Canestraro B.K. collector number, ! Engels, M.E. collector number).

Regarding life forms (Tab. 1), 63 species (48.9%) are terrestrial herbs, 23 epiphytes (17.9%), 14 lithophytes (10.9%), six terrestrial arborescents (4.6%), two aquatics (1.5%) and 21 generalists (16.3%). Considering accidental, facultative and mandatory epiphytes, 33 species were recorded (25.5%). Pecluma singeri (de la Sota) M.G.Price was the only species growing as hemiepiphyte, however it is also considered a terrestrial herb.

In regard to the vegetation type (Tab. 1), 79 species (61.2%) were growing in woody forests, 32 (24.8%) in herbaceous-shrubby environments, 16 (12.4%) in both vegetation formation and two (1.5%) in aquatic vegetation. The richest phytophysiognomies with non-overlapping species were the secondary forests with 19 species (14.7%), followed by gallery forests (18 spp. - 13.9%), human disturbed environments (15 spp. - 11.6%), riparian forests (12 spp. - 9.3), litholic grassland (5 spp. - 3.8%), swamp areas and wet grassland (4 spp. - 3.1% each), alagados reservoir (2 spp. - 1.5%) and dry grassland with one species (0.7%). Riparian slabstone environment was the only phytophysiognomy without exclusive species. One the other hand, 49 species (37.9%) were found in two or more phytophysiognomies, highlighting that the vast majority of species of ferns and lycophytes in “Campos Gerais” are not restricted to one type of environment.

Concerning the geographical distribution of the species (Tab. 1), most of them were Neotropical (48 species - 37.3%), followed by South American species (36 spp. - 27.9%), endemic to Brazil (20 spp. - 15.5%), Pantropical (16 spp. - 12.4%) and 6 (4.6%) are endemic to southern/southeastern Brazil.

Discussion

It was found in Carambeí 29% of the total species cited for the Paraná state, which presents 444 species of ferns and lycophytes (Labiak 2014bLabiak PH (2014b) Samambaias e licófitas. In: Kaehler M, Goldenberg R, Labiak PH, Ribas OS, Vieira AOS & Hatschbach GG (eds.) Plantas vasculares do Paraná. Ed. UFPR, Curitiba. Pp. 45-55.). The number of species is consistent with other studies conducted in the “Campos Gerais” region (Schwartsburd and Labiak 2007Schwartsburd PB, Labiak PH & Salino A (2007) A new species of Ctenitis (Dryopteridaceae) from southern Brazil. Brittonia 59: 29-32.; Michelon & Labiak 2013Michelon C & Labiak PH (2013) Samambaias e licófitas do Parque Estadual do Guartelá, PR, Brasil. Hoehnea 40: 191-204.), as well as the most representative families and genera (Tab. 2). The richest fern families found in Carambeí agree with several studies performed in the Atlantic Rain Forest, which constantly cite the following as the most diverse families, regardless of the order, Polypodiaceae, Thelypteridaceae, Pteridaceae, Dryopteridaceae, Aspleniaceae, Blechnaceae and Hymenophyllaceae (see Salino 1996Salino A (1996) Levantamento das pteridófitas da Serra do Cuscuzeiro, Analândia, SP, Brasil. Revista Brasileira de Botânica 19: 173-178.; Salino & Joly 2001Salino A & Joly CA (2001) Pteridophytes of three remnants of gallery forests in the Jacaré-Pepira river basin, São Paulo state, Brazil. Boletim do Herbário Ezechias Paulo Heringer 8: 5-15.; Mynssen & Windisch 2004Mynssen CM & Windisch PG (2004) Pteridófitas da Reserva Rio das Pedras, Mangaratiba, RJ, Brasil. Rodriguesia 55: 125-156.; Salino et al. 2005Salino A, Silva SM, Dittrich VAO & Britez RM (2005) Flora pteridofítica. In: Marques MCM & Britez RC (eds.) História natural e conservação da Ilha do Mel. Ed. UFPR, Curitiba. Pp. 85-101.; Nóbrega & Prado 2008Nóbrega GA & Prado J (2008) Pteridófitas da vegetação nativa do Jardim Botânico Municipal de Bauru, estado de São Paulo, Brasil. Hoehnea 35: 7-55.; Gasper & Savegnani 2010Gasper AL & Sevegnani L (2010) Lycophyta e samambaias do Parque Nacional da Serra do Itajaí, Vale do Itajaí, SC, Brasil. Hoehenea 37: 755-767.; Matos et al. 2010Matos FB, Amorim AM & Labiak PH (2010) The ferns and lycophytes of a montane tropical forest in southern Bahia, Brazil. Journal of the Botanical Research Institute of Texas 4: 333-346.; Lautert et al. 2015Lautert M, Temponi LG, Viveros RS & Salino A (2015) Lycophytes and ferns composition of Atlantic Forest conservation units in western Paraná with comparisons with other areas in southern Brazil. Acta Botanica Brasilica 29: 499-508.; Mazziero & Nonato 2015Mazziero FFF, Labiak PH & Paciencia M (2015) Checklist of ferns and lycophytes from the Parque Estadual Turístico do Alto Ribeira, Iporanga, São Paulo, Brazil. Check List 11: 1791.; Mazziero et al. 2015Mazziero FFF & Nonato FR (2015) Ferns and lycophytes from Jaú, São Paulo, Brazil. Check List 11: 1798.). It is also true for other Brazilian biomes; Prado et al. (2015)Prado J, Sylvestre LS, Labiak PH, Windisch PG, Salino A, Barros ICL, Hirai RY, Almeida TE, Santiago ACP, Kieling-Rubio MA, Pereira AFN, Øllgaard B, Ramos CG, Mickel JT, Dittrich VAO, Mynssen CM, Schwartsburd PB, Condack JPS, Pereira JBS & Matos FB (2015) Diversity of ferns and lycophytes in Brazil. Rodriguésia 66: 1073-1083. suggest that these families are the most diverse in Brazil and as well as Tryon & Tryon (1982)Tryon RM & Tryon AF (1982). Ferns and allied plants, with special reference to tropical America. Springer-Verlag, New York. 857p. in the Neotropics. Among the three families of lycophytes cited for Brazil (Prado et al. 2015Prado J, Sylvestre LS, Labiak PH, Windisch PG, Salino A, Barros ICL, Hirai RY, Almeida TE, Santiago ACP, Kieling-Rubio MA, Pereira AFN, Øllgaard B, Ramos CG, Mickel JT, Dittrich VAO, Mynssen CM, Schwartsburd PB, Condack JPS, Pereira JBS & Matos FB (2015) Diversity of ferns and lycophytes in Brazil. Rodriguésia 66: 1073-1083.), we recorded two in Carambeí (Lycopodiaceae and Selaginellaceae). These families are the most diverse in Brazil (Prado et al. 2015Prado J, Sylvestre LS, Labiak PH, Windisch PG, Salino A, Barros ICL, Hirai RY, Almeida TE, Santiago ACP, Kieling-Rubio MA, Pereira AFN, Øllgaard B, Ramos CG, Mickel JT, Dittrich VAO, Mynssen CM, Schwartsburd PB, Condack JPS, Pereira JBS & Matos FB (2015) Diversity of ferns and lycophytes in Brazil. Rodriguésia 66: 1073-1083.) and are frequently cited as the richest in several studies (e.g., Salino & Almeida 2008Salino A & Almeida TE (2008) Pteridófitas do Parque Estadual do Jacupiranga, SP, Brasil. Acta Botanica Brasilica 22: 983-991.; Mazziero et al. 2015Mazziero FFF & Nonato FR (2015) Ferns and lycophytes from Jaú, São Paulo, Brazil. Check List 11: 1798.).

Table 2
Comparison between ferns and lycophytes floras in the "Campos Gerais" region. The comparison areas are represented by Guartelá = Guartelá State Park (Michelon and Labiak 2013Michelon C & Labiak PH (2013) Samambaias e licófitas do Parque Estadual do Guartelá, PR, Brasil. Hoehnea 40: 191-204.), Vila Velha = Vila Velha State Park (Schwartsburd and Labiak 2007Schwartsburd PB & Labiak PH (2007) Pteridófitas do Parque Estadual de Vila Velha, Ponta Grossa, Paraná, Brasil. Hoehnea 34: 159-209.), Klabin = Klabin Ecological Park (Sakagami 2006Sakagami CR (2006) Pteridófitas do Parque Ecológico da Klabin, Telêmaco Borba, Paraná, Brasil. Dissertação de Mestrado, Universidade Federal do Paraná, Curitiba. 212p.). Families = the three richest families, Genera = the most representative genera, Epiphytes = number of species, Endemism = percentage of species endemic to Brazil.

Among the genera, Asplenium and Amauropelta were the richest followed by Doryopteris, Hymenophyllum and Pecluma, all of them have shown an expressive richness in the “Campos Gerais” (Tab. 2) (Schwartsburd & Labiak 2007Schwartsburd PB, Labiak PH & Salino A (2007) A new species of Ctenitis (Dryopteridaceae) from southern Brazil. Brittonia 59: 29-32.; Michelon & Labiak 2013Michelon C & Labiak PH (2013) Samambaias e licófitas do Parque Estadual do Guartelá, PR, Brasil. Hoehnea 40: 191-204.). The combination of those genera as the most expressive shows a tendency for the ferns’ diversity in Paraná’s Second Plateau vegetation (Tab. 2). In the Atlantic rain forest, Asplenium and Amauropelta usually figure as the richest genera, however this is not true for Doryopteris, Hymenophyllum and Pecluma. Blechnum, as traditionally treated, also appears as one of the most relevant genus in “Campos Gerais”, however, a recent taxonomic revision suggest their segregation in several genera (Gasper et al. 2016Gasper AL, Dittrich VAO, Smith AR & Salino A (2016) A classification for Blechnaceae (Polypodiales: Polypodiopsida): new genera, resurrected names, and combinations. Phytotaxa 275: 191-227.). Thus, now at least six genera are found in “Campos Gerais”: Austroblechnum, Blechnum, Lomariocycas, Lomaridium, Neoblechnum and Parablechnum. The same way Thelypteris, now, is divided in at least five genera in “Campos Gerais”: Amauropelta, Christella, Cyclossorus, Goniopteris and Steiropteris (Almeida et al. 2015Almeida TE, Hennequin S, Schneider H, Smith AR, Batista JAN, Ramalho AJ, Proite K & Salino A (2015) Towards a phylogenetic generic classification of Thelypteridaceae: Additional sampling suggests alteration of neotropical taxa and further study of paleotropical genera. Molecular Phylogenetics and Evolution 94: 688-700.; Salino et al. 2015Salino A, Almeida TE & Smith AR (2015) New combinations in Neotropical Thelypteridaceae. Phytokey 57: 11-50.).

Two possible hybrids were found in study area. The first one belongs to genus Anemia (Anemia sp. in Tab. 1) and the second belongs to genus Blechnum (Blechnum sp. in Tab. 1). In the first case, the specimen was found growing in a slope among Anemia tomentosa (Sav.) Sw. (subgenus Captophyllum) and Anemia phyllitidis (L.) Sw. (subgenus Anemia) and presents morphological characters of both possible parental species. It’s probably product of hybridization, a common process in the genus Anemia, but rare between subgenera (Mickel 1962Mickel JT (1962) A monographic study of the fern genus Anemia, subgenus Captophyllum. Iowa State Journal of Science 4: 349-382., 1982Mickel JT (1982) The genus Anemia (Schizaeaceae) in Mexico. Brittonia 34: 388-413.). The second case of hybridizations is Blechnum sp. which presents intermediate characters between the monomorphic species of Blechnum found in this study, such as B. austrobrasilianum and B. polypodioides. Hybridization is also common in Blechnum (Moran 1995bMoran RC (1995b) Blechnaceae. In: Davidse G, Sousa M & Knapp S (eds.) Flora Mesoamericana. Vol. 1. Universidad Nacional Autónoma do México, Ciudad de México. Pp. 325-333.), and well documented in the “Campos Gerais” region (Michelon and Labiak 2013Michelon C & Labiak PH (2013) Samambaias e licófitas do Parque Estadual do Guartelá, PR, Brasil. Hoehnea 40: 191-204.).

Terrestrial and epiphyte species are the most diverse guilds of life forms in ferns and lycophytes in Brazil (Prado et al. 2015Prado J, Sylvestre LS, Labiak PH, Windisch PG, Salino A, Barros ICL, Hirai RY, Almeida TE, Santiago ACP, Kieling-Rubio MA, Pereira AFN, Øllgaard B, Ramos CG, Mickel JT, Dittrich VAO, Mynssen CM, Schwartsburd PB, Condack JPS, Pereira JBS & Matos FB (2015) Diversity of ferns and lycophytes in Brazil. Rodriguésia 66: 1073-1083.), which agrees with our result for Carambeí. These guilds of life forms are cited in studies through Brazil independently of the biome (Mynssen & Windisch 2004Mynssen CM & Windisch PG (2004) Pteridófitas da Reserva Rio das Pedras, Mangaratiba, RJ, Brasil. Rodriguesia 55: 125-156.; Salino et al. 2005Salino A, Silva SM, Dittrich VAO & Britez RM (2005) Flora pteridofítica. In: Marques MCM & Britez RC (eds.) História natural e conservação da Ilha do Mel. Ed. UFPR, Curitiba. Pp. 85-101.; Xavier & Barros 2005Xavier SRS & Barros ICL (2005) Pteridoflora e seus aspectos ecológicos ocorrentes no Parque Ecológico João Vasconselos Sobrinho, Caruaru, Pernambuco, Brasil. Acta Botanica Brasilica 19: 775-781.; Nóbrega & Prado 2008Nóbrega GA & Prado J (2008) Pteridófitas da vegetação nativa do Jardim Botânico Municipal de Bauru, estado de São Paulo, Brasil. Hoehnea 35: 7-55.; Zuquim et al. 2008Zuquim G, Costa FRC, Prado J & Tuomisto H (2008) Guia de samambaias e licófitas da REBIO Uatumã, Amazônia central. Átemma Design Editorial, Manaus. 316p.; Gasper & Sevegnani 2010Gasper AL & Sevegnani L (2010) Lycophyta e samambaias do Parque Nacional da Serra do Itajaí, Vale do Itajaí, SC, Brasil. Hoehenea 37: 755-767.; Matos et al. 2010Matos FB, Amorim AM & Labiak PH (2010) The ferns and lycophytes of a montane tropical forest in southern Bahia, Brazil. Journal of the Botanical Research Institute of Texas 4: 333-346.; Fernandes et al. 2012Fernandes RS, Maciel S & Pietrobom MR (2012) Licófitas e monilófitas das Unidades de Conservação da Usina Hidrelétrica - UHE de Tucuruí, Pará, Brasil. Hoehnea 39: 247-285.; Souza et al. 2012Souza FS, Salino A, Viana PL & Salimena FRG (2012) Pteridófitas da Serra Negra, Minas Gerais, Brasil. Acta Botanica Brasilica 26: 378-390.; Salino et al. 2013Salino A, Mota NFO & Almeida TE (2013) Lycophytes and monilophytes in Rio Preto State Park, Minas Gerais, Brazil. Acta Botanica Brasilica 27: 252-263.; Mazziero & Nonato 2015Mazziero FFF, Labiak PH & Paciencia M (2015) Checklist of ferns and lycophytes from the Parque Estadual Turístico do Alto Ribeira, Iporanga, São Paulo, Brazil. Check List 11: 1791.; Mazziero et al. 2015Mazziero FFF & Nonato FR (2015) Ferns and lycophytes from Jaú, São Paulo, Brazil. Check List 11: 1798.) the same way in Carambeí and other studies carried out in the inlands of Paraná (Tab. 2) (Schwartsburd & Labiak 2007Schwartsburd PB, Labiak PH & Salino A (2007) A new species of Ctenitis (Dryopteridaceae) from southern Brazil. Brittonia 59: 29-32.; Michelon & Labiak 2013Michelon C & Labiak PH (2013) Samambaias e licófitas do Parque Estadual do Guartelá, PR, Brasil. Hoehnea 40: 191-204.; Lautert et al. 2015Lautert M, Temponi LG, Viveros RS & Salino A (2015) Lycophytes and ferns composition of Atlantic Forest conservation units in western Paraná with comparisons with other areas in southern Brazil. Acta Botanica Brasilica 29: 499-508.). Epiphytes were the second guild in number of species in Carambeí and in the other inland regions of Paraná (Tab. 2) (Schwartsburd & Labiak 2007Schwartsburd PB & Labiak PH (2007) Pteridófitas do Parque Estadual de Vila Velha, Ponta Grossa, Paraná, Brasil. Hoehnea 34: 159-209.; Michelon & Labiak 2013Michelon C & Labiak PH (2013) Samambaias e licófitas do Parque Estadual do Guartelá, PR, Brasil. Hoehnea 40: 191-204.; Lautert et al. 2015Lautert M, Temponi LG, Viveros RS & Salino A (2015) Lycophytes and ferns composition of Atlantic Forest conservation units in western Paraná with comparisons with other areas in southern Brazil. Acta Botanica Brasilica 29: 499-508.) as well in other areas of Atlantic Rain Forest domain (Salino 1996Salino A (1996) Levantamento das pteridófitas da Serra do Cuscuzeiro, Analândia, SP, Brasil. Revista Brasileira de Botânica 19: 173-178.; Salino & Joly 2001Salino A & Joly CA (2001) Pteridophytes of three remnants of gallery forests in the Jacaré-Pepira river basin, São Paulo state, Brazil. Boletim do Herbário Ezechias Paulo Heringer 8: 5-15.; Mynssen & Windisch 2004Mynssen CM & Windisch PG (2004) Pteridófitas da Reserva Rio das Pedras, Mangaratiba, RJ, Brasil. Rodriguesia 55: 125-156.; Nóbrega & Prado 2008Nóbrega GA & Prado J (2008) Pteridófitas da vegetação nativa do Jardim Botânico Municipal de Bauru, estado de São Paulo, Brasil. Hoehnea 35: 7-55.; Gasper & Savegnani 2010Gasper AL & Sevegnani L (2010) Lycophyta e samambaias do Parque Nacional da Serra do Itajaí, Vale do Itajaí, SC, Brasil. Hoehenea 37: 755-767.; Matos et al. 2010Matos FB, Amorim AM & Labiak PH (2010) The ferns and lycophytes of a montane tropical forest in southern Bahia, Brazil. Journal of the Botanical Research Institute of Texas 4: 333-346.; Mazziero & Nonato 2015Mazziero FFF & Nonato FR (2015) Ferns and lycophytes from Jaú, São Paulo, Brazil. Check List 11: 1798.; Mazziero et al. 2015Mazziero FFF & Nonato FR (2015) Ferns and lycophytes from Jaú, São Paulo, Brazil. Check List 11: 1798.; Lautert et al. 2015Lautert M, Temponi LG, Viveros RS & Salino A (2015) Lycophytes and ferns composition of Atlantic Forest conservation units in western Paraná with comparisons with other areas in southern Brazil. Acta Botanica Brasilica 29: 499-508.).

The preponderance of woody forests as the preferential vegetation type for ferns and lycophytes is in agreement with Prado et al. (2015)Prado J, Sylvestre LS, Labiak PH, Windisch PG, Salino A, Barros ICL, Hirai RY, Almeida TE, Santiago ACP, Kieling-Rubio MA, Pereira AFN, Øllgaard B, Ramos CG, Mickel JT, Dittrich VAO, Mynssen CM, Schwartsburd PB, Condack JPS, Pereira JBS & Matos FB (2015) Diversity of ferns and lycophytes in Brazil. Rodriguésia 66: 1073-1083. that suggest the Ombrophilous Forests are the most diverse vegetation type, especially in the Atlantic Rain Forest domain, which is corroborated by several studies in these areas, for example Salino & Almeida (2008)Salino A & Almeida TE (2008) Pteridófitas do Parque Estadual do Jacupiranga, SP, Brasil. Acta Botanica Brasilica 22: 983-991., Matos et al. (2010)Matos FB, Amorim AM & Labiak PH (2010) The ferns and lycophytes of a montane tropical forest in southern Bahia, Brazil. Journal of the Botanical Research Institute of Texas 4: 333-346., Souza et al. (2012)Souza FS, Salino A, Viana PL & Salimena FRG (2012) Pteridófitas da Serra Negra, Minas Gerais, Brasil. Acta Botanica Brasilica 26: 378-390. and Mazziero et al. (2015)Mazziero FFF, Labiak PH & Paciencia M (2015) Checklist of ferns and lycophytes from the Parque Estadual Turístico do Alto Ribeira, Iporanga, São Paulo, Brazil. Check List 11: 1791.. Secondary forests areas are the most relevant phytophysiognomies by harboring a high richness of exclusive and overlapping species. This could be related to the well preserved state of these areas and also to the fact that secondary forests provide great diversity of niches for these species. Moreover, human disturbed environments also showed high richness of exclusive and overlapping species and it is probably related to the species’s preference to open and disturbed areas, for instance: Amauropelta, Dicranopteris, Pteridium and Sticherus (Salino & Semir 2002Salino A & Semir J (2002) Thelypteridaceae (Polypodiophyta) do estado de São Paulo: Macrothelypteris e Thelypteris subgêneros Cyclosorus e Steiropteris. Lundiana 3: 9-27.; 2004Salino A & Semir J (2004) Thelypteris subg. Amauropelta (Kunze) A.R. Sm. (Thelypteridaceae - Pteridophyta) no estado de São Paulo, Brasil. Lundiana 5: 83-112.; Melthreter et al. 2010Melthreter K, Walker LR & Sharpe JM (2010) Fern ecology. Cambridge University Press, Cambridge. 460p.). An interesting fact of Carambeí ferns and lycophytes flora is the low number of species in swamp areas (Tab. 1). Generally these areas shelter a high number of species compared with other phytophysiognomies in the inlands as seen in some studies in São Paulo state by Salino (1996)Salino A (1996) Levantamento das pteridófitas da Serra do Cuscuzeiro, Analândia, SP, Brasil. Revista Brasileira de Botânica 19: 173-178., Salino & Joly (2001)Salino A & Joly CA (2001) Pteridophytes of three remnants of gallery forests in the Jacaré-Pepira river basin, São Paulo state, Brazil. Boletim do Herbário Ezechias Paulo Heringer 8: 5-15., Nóbrega & Prado (2008)Nóbrega GA & Prado J (2008) Pteridófitas da vegetação nativa do Jardim Botânico Municipal de Bauru, estado de São Paulo, Brasil. Hoehnea 35: 7-55. and Mazziero & Nonato (2015)Mazziero FFF & Nonato FR (2015) Ferns and lycophytes from Jaú, São Paulo, Brazil. Check List 11: 1798..

Species with wide geographical distribution are generally well represented in Brazilian surveys (Athayde-Filho et al. 2003Athayde Filho FP, Pereira VS, Smidt EC & Nonato FR (2003) Pteridófitas do Parque Estadual da Ilha Anchieta (PEIA). Bradea 9: 55-66.; Costa & Pietrobom 2007Costa JM & Pietrobom MR (2007) Pteridófitas (Lycophyta e Monylophyta) da Ilha de Mosqueiro, município de Belém, estado do Pará, Brasil. Boletim do Museu Paraense Emílio Goeldi 2: 45-55., 2010Costa JM & Pietrobom MR (2010) Samambaias e licófitas do Parque Ecológico de Gunma, município de Santa Bárbara do Pará, estado do Pará, Brasil. Rodriguésia 61: 223-232.; Melo & Salino 2007Melo LCN & Salino A (2007) Pteridófitas em fragmentos florestais na APA Fernão Dias, Minas Gerais, Brasil. Rodriguésia 58: 207-220.; Salino & Almeida 2008Salino A & Almeida TE (2008) Pteridófitas do Parque Estadual do Jacupiranga, SP, Brasil. Acta Botanica Brasilica 22: 983-991.; Zuquim et al. 2008Zuquim G, Costa FRC, Prado J & Tuomisto H (2008) Guia de samambaias e licófitas da REBIO Uatumã, Amazônia central. Átemma Design Editorial, Manaus. 316p.; Gasper & Savegnani 2010Gasper AL & Sevegnani L (2010) Lycophyta e samambaias do Parque Nacional da Serra do Itajaí, Vale do Itajaí, SC, Brasil. Hoehenea 37: 755-767.; Matos et al. 2010Matos FB, Amorim AM & Labiak PH (2010) The ferns and lycophytes of a montane tropical forest in southern Bahia, Brazil. Journal of the Botanical Research Institute of Texas 4: 333-346.; Fernandes et al. 2012Fernandes RS, Maciel S & Pietrobom MR (2012) Licófitas e monilófitas das Unidades de Conservação da Usina Hidrelétrica - UHE de Tucuruí, Pará, Brasil. Hoehnea 39: 247-285.; Macedo et al. 2013Macedo TS, Góes Neto A & Nonato FR (2013) Análise florística e fitogeografia das samambaias e licófitas de um fragmento de Mata Atlântica na Serra da Jibóia, Santa Terezinha, Bahia, Brasil. Rodriguésia 64: 561-572.; Mazziero et al. 2015Mazziero FFF, Labiak PH & Paciencia M (2015) Checklist of ferns and lycophytes from the Parque Estadual Turístico do Alto Ribeira, Iporanga, São Paulo, Brazil. Check List 11: 1791.) as observed in this study and in other areas of inland of Paraná (Tab. 2) (Schwartsburd & Labiak 2007Schwartsburd PB, Labiak PH & Salino A (2007) A new species of Ctenitis (Dryopteridaceae) from southern Brazil. Brittonia 59: 29-32.; Michelon & Labiak 2013Michelon C & Labiak PH (2013) Samambaias e licófitas do Parque Estadual do Guartelá, PR, Brasil. Hoehnea 40: 191-204.). We found four exotics and sub-spontaneous species in Brazil (Deparia petersenii, Macrothelypteris torresiana, Pteris vittata and Christella dentata). They have a broad geographic distribution (Prado et al. 2015Prado J, Sylvestre LS, Labiak PH, Windisch PG, Salino A, Barros ICL, Hirai RY, Almeida TE, Santiago ACP, Kieling-Rubio MA, Pereira AFN, Øllgaard B, Ramos CG, Mickel JT, Dittrich VAO, Mynssen CM, Schwartsburd PB, Condack JPS, Pereira JBS & Matos FB (2015) Diversity of ferns and lycophytes in Brazil. Rodriguésia 66: 1073-1083.) and generally occur in disturbed places in the studied areas (Salino & Almeida 2008Salino A & Almeida TE (2008) Pteridófitas do Parque Estadual do Jacupiranga, SP, Brasil. Acta Botanica Brasilica 22: 983-991.; Mazziero et al. 2015Mazziero FFF, Labiak PH & Paciencia M (2015) Checklist of ferns and lycophytes from the Parque Estadual Turístico do Alto Ribeira, Iporanga, São Paulo, Brazil. Check List 11: 1791.). In the other hand, some species could be interpreted as rare in this region of Paraná, it is the case of Arachniodes denticulata, Cyathea villosa, Phlegmariurus flexibilis and Phlegmariurus heterocarpon, all with few records in “Campos Gerais”. Tryonia areniticola is endemic to the sandstones outcrops in Paraná and São Paulo states (Schwartsburd & Labiak 2008Schwartsburd PB & Labiak PH (2008) Eriosorus areniticola (Pteridaceae), a new species from Brazil. American Fern Journal 98: 160-163.) and Dicksonia sellowiana is an endangered species in Brazil (Santiago et al. 2013Santiago ACP, Mynssen CM, Maurenza D, Penedo TSA & Sfair JC (2013) Dickoniaceae. In: Martinelli G & Moraes MA (eds.) Livro vermelho da flora do Brasil. Centro Nacional de Conservação da Flora, Rio de Janeiro. Pp. 475-476.).

Our study shows the importance of preserving the remnants of vegetation in Carambeí municipality, included in the “Campos Gerais”, even for the small ones, given the relevant number of species as well as by the presence of endangered and rare species of ferns and lycophytes.

  • Editora de área: Dra. Lana Sylvestre

Acknowledgments

The authors are grateful to reviewers and PhD. Fabiana R. Nonato for help in reviewing the English. We also thank PhD. Lana S. Sylvestre by the determination of Asplenium jucundum.

References

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Publication Dates

  • Publication in this collection
    Apr-Jun 2018

History

  • Received
    24 Nov 2016
  • Accepted
    11 Aug 2017
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