Abstract:

The Porto Ferreira State Park (PFSP) is located in the State of São Paulo southeastern Brazil, in an intriguing transitional area between the Atlantic Forest and Cerrado - both hotspots of biodiversity - represented mainly by the cerradão (CER), and the seasonal semideciduous forest (SSF), with its alluvial variation vegetation type (riparian forest - RP). Ecotonal areas play an important role in providing ecological and phytogeographic knowledge regarding the flora and vegetation of this region. Despite various studies on the PFSP, knowledge of this region remains fragmented. In this study, we aim to conduct an updated checklist of the PFSP vascular flora, including a compilation of all the studies conducted in this protected area, plus field work carried out by the authors from 2014 to 2017. In addition, given its ecotonal characteristics, we completed a floristic similarity analysis between the PFSP and other floristic surveys that examined the same vegetation types present in this study, to gain a better understanding of their phytogeographic relationships. Overall, 684 species, belonging to 387 genera and 107 families, were recorded. The SSF presented the richest vegetation type (478 species), followed by the CER (418) and the RP (231). The most diverse families were Fabaceae (64 species), Myrtaceae (41), Orchidaceae (39), Rubiaceae (37), Asteraceae (35), Bignoniaceae (26) and Malvaceae (20). Moreover, eight threatened species, at regional and national levels, were found. To date, 412 species have been added to the floristic list produced for the PFSP. The life forms with the highest number of species were trees (286 species), herbs (176) plus shrubs and subshrubs (123). Our research findings indicate floristic patterns with higher levels of similarity among species in geographical proximity, including those in ecotonal areas encompassing different vegetation types. These results rank the PFSP among some of the most species-rich conservation units with seasonal climates, and therefore is of great importance for plant conservation in the southeast of Brazil.

Keywords:
Atlantic Forest; Cerrado; floristic; protected area; seasonal semideciduous forest

Resumo:

O Parque Estadual de Porto Ferreira (PEPF) está localizado no estado de São Paulo, sudeste do Brasil, em uma intrigante área de contato entre a Mata Atlântica e o Cerrado - ambos hotspots de biodiversidade -, representada principalmente pelo cerradão (CER), a floresta estacional semidecidual (FES) e pela variação dessa última, com influência aluvial (FESA). As áreas ecotonais desempenham um papel importante no conhecimento ecológico e fitogeográfico da flora e da vegetação. Mesmo com muitos estudos realizados no PEPF, as informações encontram-se dispersas. Dessa forma, produzimos um checklist atualizado da flora vascular, resultante da compilação de vouchers, estudos realizados no PEPF e de esforços de coleta realizados pelos autores entre os anos de 2014 e 2017. Além disso, para entender suas relações fitogeográficas, realizamos uma análise de similaridade florística entre o PEPF e outros levantamentos realizadas nas mesmas fitofisionomias da área estudada. Registramos 684 espécies pertencentes a 387 gêneros e 107 famílias. A FES foi a fitofisionomia mais rica (478 espécies), seguida pelo CER (418) e a FESA (231). As famílias mais ricas foram Fabaceae (64 espécies), Myrtaceae (41), Orchidaceae (39), Rubiaceae (37), Asteraceae (35), Bignoniaceae (26) e Malvaceae (20). Oito espécies ameaçadas em âmbito regional e nacional foram encontradas. Desde a última lista florística produzida para o PEPF, 412 espécies foram adicionadas. As formas de vida com maiores números de espécies foram: árvores (286 espécies), ervas (176) e arbustos mais subarbustos (123). A análise de similaridade corroborou o padrão de que áreas mais próximas geograficamente são mais similares entre si, e esse padrão foi encontrado, também, para áreas ecotonais envolvendo diferentes fitofisionomias. Os resultados encontrados colocam o PEPF entre as unidades de conservação com maior número de espécies vegetais sob o domínio de clima sazonal, possuindo, portanto, grande importância para a conservação de espécies de plantas no Sudeste do Brasil.

Palavras-chave:
Mata Atlântica; Cerrado; florística; unidade de conservação; floresta estacional semidecidual

Introduction

Brazil is home to one of the richest plant diversities in the world, with more than 34,000 species of vascular plants (BFG, 2021BFG. 2021. Flora do Brasil 2020. Jardim Botânico do Rio de Janeiro, Rio de Janeiro, p.1-28.). This great diversity is mainly due to Brazil’s vast geographical area, its heterogeneous topography, as well as its climate, and biogeographical features (Fiaschi & Pirani, 2009FIASCHI, P. & PIRANI, J.R. 2009. Review of plant biogeographic studies in Brazil. J. Syst. Evol. 47(5):477-496.). Traditionally, it has been divided into major landscapes and macroecological ecoregions (Ab’Sáber, 1970AB’SÁBER, A.N. 1970. Províncias geológicas e domínios morfoclimáticos no Brasil. IGEOG/USP, São Paulo., 2003AB’SÁBER, A.N. 2003. Os domínios de natureza do Brasil: Potencialidades paisagísticas. Ateliê Editorial, São Paulo.). Among the biogeographical units recognized in Brazil, the Cerrado and the Atlantic Forest are classified as biodiversity hotspots for global conservation, hosting an overwhelming number of endemic species threatened by anthropic actions (Myers et al., 2000MYERS, N., MITTERMEIER, R.A., MITTERMEIER, C.G., FONSECA, G.A.B. & KENT, J. 2000. Biodiversity hotspots for conservation priorities. Nature. 403:852-858.; Mittermeier et al., 2004MITTERMEIER, R.A., ROBLES-GIL, P., HOFFMANN, M., PILGRIM, J., BROOKS, T., MITTERMEIER, C.G., LAMOREUX, J. & FONSECA, G.A.B. 2004. Hotspots revisited: Earth’s biologically richest and most endangered terrestrial ecoregions. EMEX Agrupacíon Sierra Madre, Cidade do México.).

The biogeographic province, known as the Cerrado, mainly spans over central Brazil, with its southern limits reaching the northern part of the State of Paraná, the northern region of Argentina, and eastern Bolivia (Morrone, 2017MORRONE, J.J. 2017. Neotropical biogeography: regionalization and evolution. CRC Press, Boca Raton.; Velazco et al., 2018VELAZCO, S.J.E., GALVÃO, F., KELLER, H.A. & BEDRIJ, N.A. 2018. Cerrados in Argentina? Structure, diversity and biogeography of the woody component of a savanna in the Misiones Province. Rodriguésia. 69(2):335-349.). It comprises a set of vegetation types, ranging from grasslands with herbaceous plants dominating the landscape (campo limpo), to savannas (campo sujo and cerrado sensu stricto) and forests (e.g. cerradão and gallery forests) (Coutinho, 2006COUTINHO, L.M. 2006. O conceito de bioma. Acta Bot. Bras. 20(1):13-23.; Batalha, 2011BATALHA, M.A. 2011. O cerrado não é um bioma. Biota Neotropica. 11(1):21-24. https://www.biotaneotropica.org.br/v11n1/en/abstract?inventory+bn00111012011 (last access in 01/03/2020)
https://www.biotaneotropica.org.br/v11n1... ). The Cerrado is the second largest phytogeographical unit of Brazil, spanning 2 million km² or 23.9% of the national territory (IBGE, 2004IBGE. 2004. Mapa de Biomas do Brasil. Diretoria de Geociências, Rio de Janeiro.). The main threats to the Cerrado vegetation are the expansion of intensive agriculture activity, and raising livestock (Ratter et al., 2006RATTER, J.A.; BRIDGEWATER, S.; RIBEIRO, J.F. 2006. Biodiversity patterns of the woody vegetation of the Brazilian Cerrado. In: PENNINGTON, T.; LEWIS GWILYM, P.; RATTER, J.A. (Org.). Neotropical Savannas and dry forests: diversity, biogeography and conservation. Boca Raton: Taylor & Francis. p.31-66.; Klink & Moreira, 2002KLINK, C.A. & MOREIRA, A.G. 2002. Past and current human occupation, and land use. In: The Cerrados of Brazil, OLIVEIRA, P.S. & MARQUIS, R.J. (eds). Columbia University Press. New York. USA. p 69.). Nevertheless, the Cerrado is considered one of the world’s richest savanna vegetation (Klink & Machado, 2005KLINK, C.A. & MACHADO, R.B. 2005. A conservação do Cerrado Brasileiro. Megadiversidade. 1(1):147-155.; Simon et al. 2009SIMON, M.F., GRETHER, R., QUEIROZ, L.P., SKEMA, C., PENNINGTON, R.T. & HUGHES, C.E. 2009. Recent assembly of the Cerrado, a neotropical plant diversity hotspot, by in situ evolution of adaptations to fire. PNAS. 106(48):20359-20364.), containing more than 12,000 species of vascular plants (BFG, 2015BFG. 2015. Growing knowledge: an overview of Seed Plant diversity in Brazil. Rodriguésia. 66(4):1085-1113.; Flora do Brasil, 2020FLORA DO BRASIL. 2020. Jardim Botânico do Rio de Janeiro. http://floradobrasil.jbrj.gov.br/(last access in 12/05/2020)
http://floradobrasil.jbrj.gov.br/... ). In the State of São Paulo the Cerrado occurs mainly in the central region, but extends to eastern portions of the State as well, with enclaves in the Atlantic Forest matrix (Baitello et al., 2013BAITELLO, J.B., AGUIAR, O.T., PASTORE, J.A. & ARZOLLA, F.A.R.D.P.A. 2013. Parque Estadual do Juquery: refúgio de Cerrado no Domínio Atlântico. IF. Ser. Regist. 50:1-46).

The Atlantic Forest covers nearly 1.1 million km² (IBGE, 2004IBGE. 2004. Mapa de Biomas do Brasil. Diretoria de Geociências, Rio de Janeiro.), distributed over a broad latitudinal, ranging from the State of Piauí to Rio Grande do Sul (SOS Mata Atlântica, 2016SOS MATA ATLÂNTICA. 2016. Atlas dos remanescentes florestais da Mata Atlântica no período 2014-2015. Fundação SOS Mata Atlântica, São Paulo. http://mapas.sosma.org.br/ (last access in 15/08/2019)
http://mapas.sosma.org.br/... ), and reaching inland to northern Argentina and eastern Paraguay (Spichiger et al., 2006SPICHIGER, R., BISE, B., CALENGE, C. & CHATELAIN, C. 2006. Biogeography of the forest of the Paraguay-Parana Basin. In Neotropical savannas and seasonally dry forests plant diversity, biogeography, and conservation (R.T. Pennington & J.A. Ratter, eds). Taylor & Francis, Boca Raton, p.193-209.; Fiaschi & Pirani, 2009FIASCHI, P. & PIRANI, J.R. 2009. Review of plant biogeographic studies in Brazil. J. Syst. Evol. 47(5):477-496.). Brazil’s Atlantic Forest hosts an overwhelming species richness of vascular plants (BFG 2015BFG. 2015. Growing knowledge: an overview of Seed Plant diversity in Brazil. Rodriguésia. 66(4):1085-1113.; Flora do Brasil, 2020FLORA DO BRASIL. 2020. Jardim Botânico do Rio de Janeiro. http://floradobrasil.jbrj.gov.br/(last access in 12/05/2020)
http://floradobrasil.jbrj.gov.br/... ), being one of the world’s top five species-rich areas (Barthlott et al., 2005BARTHLOTT, W., MUTKE, J., RAFIQPOOR, M.D., KIER, G. & KREFT, H. 2005. Global centers of vascular plant diversity. Nova. Act. Lc. 92:61-83.). Traditionally, the Atlantic Forest has been considered a single biogeographical unit with subdivisions (e.g.Morellato & Haddad, 2000MORELLATO, L.P.C. & HADDAD, C.F. 2000. Introduction: The Brazilian Atlantic Forest. Biotropica. 32(4b):786-792.). However, more than distinct vegetation types, singular biogeographical histories of the biota have been suggested for different parts of the Atlantic Forest (Vieira et al., 2015VIEIRA, L.T.A., POLISEL R.T., IVANAUSKAS, N.M., SHEPHERD, G.J., WAECHTER, J.L., YAMAMOTO, K. & MARTINS, F.R. 2015. Geographical patterns of terrestrial herbs: a new component in planning the conservation of the Brazilian Atlantic Forest. Biodivers. Conserv. 24(9):2181-2198.; Morrone 2014MORRONE, J.J. 2014. Biogeographical regionalisation of the Neotropical region. Zootaxa. 3782:1-110., 2017MORRONE, J.J. 2017. Neotropical biogeography: regionalization and evolution. CRC Press, Boca Raton.; Eisenlohr & Oliveira-Filho, 2015EISENLOHR, P.V. & OLIVEIRA-FILHO, A.T. 2015. Revisiting Patterns of Tree Species Composition and their Driving Forces in the Atlantic Forests of Southeastern Brazil. Biotropica. 47(6):689-701.) justifying its division in three provinces (Morrone 2014MORRONE, J.J. 2014. Biogeographical regionalisation of the Neotropical region. Zootaxa. 3782:1-110., 2017MORRONE, J.J. 2017. Neotropical biogeography: regionalization and evolution. CRC Press, Boca Raton.). Among these biogeographical provinces, the inland and seasonal vegetation type is represented by the Paraná Forest province (Morrone, 2017MORRONE, J.J. 2017. Neotropical biogeography: regionalization and evolution. CRC Press, Boca Raton.; Zanotti et al., 2020ZANOTTI, C.A. KELLER, H.A. & ZULOAGA, F.O. 2020. Biodiversidad de la flora vascular de la provincia de Misiones, Región Paranaense Argentina. Darwiniana. 8(1):42-291.), and it is the one most threatened by the reduction of the original area (Ribeiro et al., 2009RIBEIRO, M.C., METZGER, J.P., MARTENSEN, A.C., PONZONI, F.J. & HIROTA, M.M. 2009. The Brazilian Atlantic forest: How much is left, and how is the remaining forest distributed? Implications for conservation. Biol. Cons. 142(6):1141-1153.). The floristic richness of the seasonal forest is considerably high (Souza et al., 2019aSOUZA, V.C., TOLEDO, C.P., SAMPAIO, D., BÍGIO, N.C., COLLETA, G.D., IVANAUSKAS, N.M. & FLORES, T.B. 2019a. Guia das plantas da mata atlântica: floresta estacional. Liana Produções Editoriais, Piracicaba.) and holds regional and typical species (Eisenlohr & Oliveira-Filho, 2015EISENLOHR, P.V. & OLIVEIRA-FILHO, A.T. 2015. Revisiting Patterns of Tree Species Composition and their Driving Forces in the Atlantic Forests of Southeastern Brazil. Biotropica. 47(6):689-701.; Vieira et al., 2015VIEIRA, L.T.A., POLISEL R.T., IVANAUSKAS, N.M., SHEPHERD, G.J., WAECHTER, J.L., YAMAMOTO, K. & MARTINS, F.R. 2015. Geographical patterns of terrestrial herbs: a new component in planning the conservation of the Brazilian Atlantic Forest. Biodivers. Conserv. 24(9):2181-2198.; Morrone 2014MORRONE, J.J. 2014. Biogeographical regionalisation of the Neotropical region. Zootaxa. 3782:1-110., 2017MORRONE, J.J. 2017. Neotropical biogeography: regionalization and evolution. CRC Press, Boca Raton.).

In the State of São Paulo, the Cerrado and Atlantic Forest mix in a complex and transitional area, in scattered patches of each or at ecotones (Durigan et al., 2012DURIGAN, G., RAMOS, V.S., IVANAUSKAS, N.M. & FRANCO, G.A.D.C. 2012. Espécies indicadoras de fitofisionomias na transição cerrado-mata atlântica no estado de São Paulo. SMA/CBRN, São Paulo.). The Cerrado forest formation is very common there, with a great richness of tree species, mainly influenced by contact with Atlantic Forest taxa (Françoso et al., 2016FRANÇOSO, R.D., HAIDAR, R.F. & MACHADO, R.B. 2016. Tree species of South America central savanna: endemism, marginal areas and the relationship with other biomes. Acta Bot. Bras. 30(1):78-86.). On the other hand, the innermost vegetation type of the Atlantic Forest is characterized by a well-marked dry season, and a characteristic composition, despite sharing several species with the rainy coastal forest and the cerradão (Eisenlohr & Oliveira-Filho, 2015EISENLOHR, P.V. & OLIVEIRA-FILHO, A.T. 2015. Revisiting Patterns of Tree Species Composition and their Driving Forces in the Atlantic Forests of Southeastern Brazil. Biotropica. 47(6):689-701.). Ecotones represent a transition between ecological communities, with a mixture of the floristic and faunistic characteristics of two different and relatively homogenous ecological community types (Allen & Starr, 1982ALLEN, T.F.H. & STARR, T.B. 1982. Hierarchy. Perspectives for ecological complexity. University of Chicago Press, Chicago.). Studies indicate that richness and abundance tend to peak at ecotonal areas (Kark & van Rensburg, 2006KARK S. & VAN RENSBURG B. J. 2006. Ecotones: marginal or central areas of transition? Isr. J. Ecol. Evol. 52:29-53.) and contribute to São Paulo’s high level of plant richness, having one of Brazil’s greatest numbers of species (BFG, 2015BFG. 2015. Growing knowledge: an overview of Seed Plant diversity in Brazil. Rodriguésia. 66(4):1085-1113.; Flora do Brasil, 2020FLORA DO BRASIL. 2020. Jardim Botânico do Rio de Janeiro. http://floradobrasil.jbrj.gov.br/(last access in 12/05/2020)
http://floradobrasil.jbrj.gov.br/... ).

Studies on the biodiversity of plant communities are considered fundamental, and one of the first steps toward the establishment of models for the preservation and conservation of ecosystems (Morellato & Leitão-Filho, 1992MORELLATTO, L.P.C. & LEITÃO-FILHO, H.F. 1992. Padrões de frutificação e dispersão na Serra do Japi. In História natural da Serra do Japi: ecologia e preservação de uma área florestal no Sudeste do Brasil (L.P.C. Morellatto, ed). Editora da UNICAMP/FAPESP, Campinas, p.112-140.; Ferreira Júnior et al., 2008FERREIRA JÚNIOR, E.V., SOARES, T.S., COSTA, M.F.F., SILVA, V.S.M. 2008. Composição, diversidade e similaridade florística de uma floresta tropical semidecídua submontana em Marcelândia - MT, Brasil. Acta Amaz. 38(4):673-680.). Given these characteristics, the PFSP, floristically, has drawn great attention. From 1984 to present, 12 studies regarding its flora have been carried out, as well as aleatory collections deposited in herbaria. These studies include climbing plants (Vargas et al., 2018VARGAS, B.C., OLIVEIRA, A.P.C., UDULUTSCH, R.G., MARCUSSO, G. M., SABINO, G.P., MELO, P.H.A., GRILLO, R.M.M., KAMIMURA, V.A. & ASSIS, M.A. 2018. Climbing plants of Porto Ferreira State Park, southeastern Brazil. Biota Neotropica. 18(2):e20170346. https://doi.org/10.1590/1676-0611-bn-2017-0346 (last access in 10/02/2020)
https://doi.org/10.1590/1676-0611-bn-201... ), ferns (Colli et al., 2003COLLI, A.M.T., SOUZA, S.A. & SILVA, R.T. 2003. Pteridófitas do Parque Estadual de Porto Ferreira (SP), Brasil. Rev. Inst. Flor. 15(11):29-35.), non-arboreal plants (Oliveira, 2012OLIVEIRA, A.P.C. 2012. Flora vascular não-arbórea do Parque Estadual de Porto Ferreira, SP, Brasil. Dissertação de Mestrado, Universidade Estadual Paulista, Rio Claro.; Osaco, 2012OSACO, M. 2012. Florística e fitossociologia do estrato arbustivo de área detransição savana-floresta no Parque Estadual de Porto Ferreira, SP. Dissertação de Mestrado, Universidade Estadual Paulista, Rio Claro.), trees (Bertoni, 1984BERTONI, J.E.A. 1984. Composição florística e estrutura fitossociológica de uma floresta de interior do Estado de São Paulo: Reserva Estadual de Porto Ferreira. Dissertação de Mestrado, Universidade Estadual de Campinas, Campinas.; Bertoni & Martins, 1987BERTONI, J.E.A. & MARTINS, F.R. 1987. Composição florística de uma floresta ripária na Reserva Estadual de Porto Ferreira, SP. Acta. Bot. Bras. 1(1):17-26.; Bertoni et al., 2001BERTONI, J.E.A., TOLEDO FILHO, D.V., LEITÃO-FILHO, H.F., FRANCO, G.A.D.C. & AGUIAR, O.T. 2001. Flora arbórea e arbustiva do cerrado do Parque Estadual de Porto Ferreira (SP). Rev. Inst. Florest. 13(2):169-188.; Sabino, 2013SABINO, G.P. 2013. Florística e Fitossociologia de uma comunidade arbórea em contato savana-floresta no Parque Estadual de Porto Ferreira, Porto Ferreira, São Paulo. Trabalho de Conclusão de Curso, Universidade Estadual Paulista, Rio Claro.; Konopczyk, 2014KONOPCZYK, R.M.G. 2014. Composição florística, estrutura e heterogeneidade ambiental da comunidade arbórea de uma várzea em Porto Ferreira, SP, Brasil. Dissertação de Mestrado, Universidade Estadual Paulista, Rio Claro.) and vascular epiphytes (Marcusso et al., 2016MARCUSSO, G.M., DICKFELDT, E.P., BERTONI, J.E.A. & MONTEIRO, R. 2016. Epífitas vasculares do Parque Estadual de Porto Ferreira, São Paulo, Brasil. Rev. Inst. Flor. 28(2):119-133.). However, this data is fragmented, and remains to be systematically organized. The Management Plan (São Paulo, 2003SÃO PAULO. 2003. Plano de manejo do Parque Estadual de Porto Ferreira, Instituto Florestal, São Paulo. https://www.infraestruturameioambiente.sp.gov.br/fundacaoflorestal/planos-de-manejo/planos-de-manejo-planos-concluidos/plano-de-manejo-pe-porto-ferreira/ (last access in 02/03/2020)
https://www.infraestruturameioambiente.s... ) represents the last overall floristic list presented for this protected area.

Thus, we compiled the checklist of the vascular flora of the PFSP, based on fieldwork, studies conducted in the area, and herbarium collections, aiming to extend and systematize the scattered floristic information on the region. This data can support restoration and conservation projects in the central region of the State of São Paulo, given that several private lands of the region present a deficit in native vegetation and must be restored in accordance with the Native Vegetation Protection Law (Tavares et al., 2019TAVARES, P.A., BRITES, A.D., SPAROVEK, G., GUIDOTTI, V., CERIGNONI, F., AGUIAR, D., METZGER, J.P.W., RODRIGUES, R.R., PINTO, L.F.G., MELLO, K. & MOLIN, P. G. 2019. Unfolding additional massive cutback effects of the Native Vegetation Protection Law on Legal Reserves, Brazil. Biota Neotropica. 19(4):e20180658. https://doi.org/10.1590/1676-0611-bn-2018-0658 (last access in 22/07/2020)
https://doi.org/10.1590/1676-0611-bn-201... ). Our work also provides a foundation on which to update the Management Plan of this important vegetation-protected area. In addition, we explored and discussed the floristic identity concerning the ecotonal condition of the PFSP, comparing our data with other floristic surveys carried out on the same vegetation types.

Material and Methods

The Porto Ferreira State Park (PFSP), known locally as “Mata do Procópio”, is located in the municipality of Porto Ferreira, of the State of São Paulo, in the southeastern region of Brazil (Figure 1). The PFSP is one of more than 50 integral protected areas in São Paulo (Instituto Florestal/Fundação Florestal, 2008INSTITUTO FLORESTAL/FUNDAÇÃO FLORESTAL. 2008. Unidades de Conservação. Arquivos Digitais - APAS Est. SP (CPLA); PESM (ITESP); Mosaico de Jacupiranga; Mosaico da Juréia; Zonas de Amortecimento conforme Planos de Manejos e Áreas Naturais Protegidas do Estado de São Paulo. Florestar Estatístico, v. 11, n. 20., Colli-Silva et al., 2016COLLI-SILVA, M., BEZERRA, T.L., FRANCO, G.A.D.C., IVANAUSKAS, N.M. & SOUZA, F.M. 2016. Registros de espécies vasculares em unidades de conservação e implicações para a lista da flora ameaçada de extinção no estado de São Paulo. Rodriguésia. 67(2):405-425.), founded in 1962, initially categorized as a State Reserve, and later (1987) becoming a State Park (São Paulo, 2003SÃO PAULO. 2003. Plano de manejo do Parque Estadual de Porto Ferreira, Instituto Florestal, São Paulo. https://www.infraestruturameioambiente.sp.gov.br/fundacaoflorestal/planos-de-manejo/planos-de-manejo-planos-concluidos/plano-de-manejo-pe-porto-ferreira/ (last access in 02/03/2020)
https://www.infraestruturameioambiente.s... ). Under the central geographical coordinates 21°51’S and 47°25’W, the park is bordered by the Mogi Guaçu river to the south, and by the SP-215 highway to the north, occupying an area of 611.55 hectares (São Paulo, 2003SÃO PAULO. 2003. Plano de manejo do Parque Estadual de Porto Ferreira, Instituto Florestal, São Paulo. https://www.infraestruturameioambiente.sp.gov.br/fundacaoflorestal/planos-de-manejo/planos-de-manejo-planos-concluidos/plano-de-manejo-pe-porto-ferreira/ (last access in 02/03/2020)
https://www.infraestruturameioambiente.s... ). The PFSP is part of the region of the Peripheral Depression of São Paulo (Depression of Mogi Guaçu) belonging to the Paraná Basin (Rossi et al., 2005ROSSI, M., MATTOS, I.F.A., COELHO, R.M., MENK, J.R.F., ROCHA, F.T., PFEIFER, R.M. & DEMARIA, I.C. 2005. Relação solos/vegetação em área natural no Parque Estadual de Porto Ferreira, São Paulo. Rev. Inst. Flor. 17(1):45-61.).

Figure 1
Location of Porto Ferreira State Park (PFSP), Porto Ferreira, São Paulo State. A. Atlantic Forest and the Cerrado in Brazil and the location of São Paulo State; B. Location of PFSP; C. PFSP satellite view, adapted from Google Earth.

According to Köeppen-Geiger classification, the climate of the region is classified as Cwa, dry winter mesothermal (Alvares et al., 2013ALVARES, C.A., STAPE, J.L., SENTELHAS, P.C., GONÇALVES, J.L.M. & SPAROVEK, G. 2013. Köppen’s climate classification map for Brazil. Meteorol. Z. 22(6):711-728.). The annual mean temperature is 22ºC, and the annual mean precipitation is 1470 mm (INMET 2019INMET. 2019. http://www.inmet.gov.br/portal/index.php?r=home2/index (last access in 12/10/2019)
http://www.inmet.gov.br/portal/index.php... ).

The vegetation of the Mogi Guaçu basin is mainly composed of Atlantic Forest, with seasonal semideciduous forest (SSF) being the most common vegetation type (Bertoni 1984BERTONI, J.E.A. 1984. Composição florística e estrutura fitossociológica de uma floresta de interior do Estado de São Paulo: Reserva Estadual de Porto Ferreira. Dissertação de Mestrado, Universidade Estadual de Campinas, Campinas.; Veloso & Góes Filho, 1982VELOSO, H.P. & GÓES FILHO, L. 1982. Fitogeografia brasileira - classificação fisionômicoecológica da vegetação neotropical. IBGE/Projeto Radambrasil, Salvador.). However, PFSP is composed of different vegetation types, presenting the forested savanna in the north (CER, with altitudes between 605-575 m), and to the south, SSF (with altitudes between 575-535 m) (Rossi et al., 2005ROSSI, M., MATTOS, I.F.A., COELHO, R.M., MENK, J.R.F., ROCHA, F.T., PFEIFER, R.M. & DEMARIA, I.C. 2005. Relação solos/vegetação em área natural no Parque Estadual de Porto Ferreira, São Paulo. Rev. Inst. Flor. 17(1):45-61.). Parallel to the Mogi Guaçu river, riparian forest is found (RP, with an average altitude of 530 m) (Bertoni & Martins, 1987BERTONI, J.E.A. & MARTINS, F.R. 1987. Composição florística de uma floresta ripária na Reserva Estadual de Porto Ferreira, SP. Acta. Bot. Bras. 1(1):17-26.; Konopczyk, 2014KONOPCZYK, R.M.G. 2014. Composição florística, estrutura e heterogeneidade ambiental da comunidade arbórea de uma várzea em Porto Ferreira, SP, Brasil. Dissertação de Mestrado, Universidade Estadual Paulista, Rio Claro.) (Figure 2). These vegetation types can be distinguished mainly by their structures, of which the CER presents a small average height and high density, while the SSF presents a greater average height and lower density (Vargas et al. 2018VARGAS, B.C., OLIVEIRA, A.P.C., UDULUTSCH, R.G., MARCUSSO, G. M., SABINO, G.P., MELO, P.H.A., GRILLO, R.M.M., KAMIMURA, V.A. & ASSIS, M.A. 2018. Climbing plants of Porto Ferreira State Park, southeastern Brazil. Biota Neotropica. 18(2):e20170346. https://doi.org/10.1590/1676-0611-bn-2017-0346 (last access in 10/02/2020)
https://doi.org/10.1590/1676-0611-bn-201... ), and finally, the RP having the lowest density and diversity (Konopczyk, 2014KONOPCZYK, R.M.G. 2014. Composição florística, estrutura e heterogeneidade ambiental da comunidade arbórea de uma várzea em Porto Ferreira, SP, Brasil. Dissertação de Mestrado, Universidade Estadual Paulista, Rio Claro.). In addition, there are ecotonal areas among them (Sabino, 2013SABINO, G.P. 2013. Florística e Fitossociologia de uma comunidade arbórea em contato savana-floresta no Parque Estadual de Porto Ferreira, Porto Ferreira, São Paulo. Trabalho de Conclusão de Curso, Universidade Estadual Paulista, Rio Claro.), as well as riverine variations (Konopczyk, 2014KONOPCZYK, R.M.G. 2014. Composição florística, estrutura e heterogeneidade ambiental da comunidade arbórea de uma várzea em Porto Ferreira, SP, Brasil. Dissertação de Mestrado, Universidade Estadual Paulista, Rio Claro.). The PFSP represents one of the few protected areas in São Paulo which comprises the contact between the Cerrado and Altantic Forest, becoming an important biotic pool of both, in a single and relatively small area. It also holds one of the largest and tallest populations of Cariniana legalis (Mart.) Kuntze and Cariniana estrellensis (Raddi) Kuntze (Lecythidaceae), with several individuals reaching up to 50 m. These trees are iconic elements of the PFSP.

Figure 2
Vegetation types of Porto Ferreira State Park (PFSP), Porto Ferreira, São Paulo State. A: map indicating the vegetation types; B: aerial view of PFSP; C: seasonal semideciduous forest; D: cerradão; E: riparian forest; F: Mogi Guaçu river (A. Adapted from Google Earth; B. PFSP photo collection; C./F. Gabriel Mendes Marcusso; D. PFSP photo collection; E. Rafael Konopczyk).

The PFSP vascular flora checklist was created from the following: specimens collected by the authors from 2014 to 2017, using the walking research method (Filgueiras et al., 1994FILGUEIRAS, T.S., NOGUEIRA, P.E., BROCHADO, A.L. & GUALALL, G.F. 1994. Caminhamento: um método expedito para levantamentos florísticos qualitativos. Cad. Geociênc. 12:39-347.); the systematization of all other lists found in surveys carried out in PFSP (Bertoni, 1984BERTONI, J.E.A. 1984. Composição florística e estrutura fitossociológica de uma floresta de interior do Estado de São Paulo: Reserva Estadual de Porto Ferreira. Dissertação de Mestrado, Universidade Estadual de Campinas, Campinas.; Bertoni & Martins, 1987BERTONI, J.E.A. & MARTINS, F.R. 1987. Composição florística de uma floresta ripária na Reserva Estadual de Porto Ferreira, SP. Acta. Bot. Bras. 1(1):17-26.; Bertoni et al., 2001BERTONI, J.E.A., TOLEDO FILHO, D.V., LEITÃO-FILHO, H.F., FRANCO, G.A.D.C. & AGUIAR, O.T. 2001. Flora arbórea e arbustiva do cerrado do Parque Estadual de Porto Ferreira (SP). Rev. Inst. Florest. 13(2):169-188.; São Paulo, 2003SÃO PAULO. 2003. Plano de manejo do Parque Estadual de Porto Ferreira, Instituto Florestal, São Paulo. https://www.infraestruturameioambiente.sp.gov.br/fundacaoflorestal/planos-de-manejo/planos-de-manejo-planos-concluidos/plano-de-manejo-pe-porto-ferreira/ (last access in 02/03/2020)
https://www.infraestruturameioambiente.s... ; Colli et al., 2003COLLI, A.M.T., SOUZA, S.A. & SILVA, R.T. 2003. Pteridófitas do Parque Estadual de Porto Ferreira (SP), Brasil. Rev. Inst. Flor. 15(11):29-35.; Oliveira, 2012OLIVEIRA, A.P.C. 2012. Flora vascular não-arbórea do Parque Estadual de Porto Ferreira, SP, Brasil. Dissertação de Mestrado, Universidade Estadual Paulista, Rio Claro.; Osaco, 2012OSACO, M. 2012. Florística e fitossociologia do estrato arbustivo de área detransição savana-floresta no Parque Estadual de Porto Ferreira, SP. Dissertação de Mestrado, Universidade Estadual Paulista, Rio Claro.; Dickfeldt et al., 2013DICKFELDT, E.P., JANDUCCI, B.Z. & SOUZA, A.S. 2013. Levantamento das espécies vegetais exóticas e experiências de manejo no Parque Estadual de Porto Ferreira, SP. Fórum Ambiental da Alta Paulista. 9(3):118-137.; Sabino, 2013SABINO, G.P. 2013. Florística e Fitossociologia de uma comunidade arbórea em contato savana-floresta no Parque Estadual de Porto Ferreira, Porto Ferreira, São Paulo. Trabalho de Conclusão de Curso, Universidade Estadual Paulista, Rio Claro.; Konopczyk, 2014KONOPCZYK, R.M.G. 2014. Composição florística, estrutura e heterogeneidade ambiental da comunidade arbórea de uma várzea em Porto Ferreira, SP, Brasil. Dissertação de Mestrado, Universidade Estadual Paulista, Rio Claro.; Marcusso et al., 2016MARCUSSO, G.M., DICKFELDT, E.P., BERTONI, J.E.A. & MONTEIRO, R. 2016. Epífitas vasculares do Parque Estadual de Porto Ferreira, São Paulo, Brasil. Rev. Inst. Flor. 28(2):119-133.; Vargas et al., 2018VARGAS, B.C., OLIVEIRA, A.P.C., UDULUTSCH, R.G., MARCUSSO, G. M., SABINO, G.P., MELO, P.H.A., GRILLO, R.M.M., KAMIMURA, V.A. & ASSIS, M.A. 2018. Climbing plants of Porto Ferreira State Park, southeastern Brazil. Biota Neotropica. 18(2):e20170346. https://doi.org/10.1590/1676-0611-bn-2017-0346 (last access in 10/02/2020)
https://doi.org/10.1590/1676-0611-bn-201... ); and the compilation of vouchers present in the Specieslink database (CRIA, 2019CRIA. 2019. SpeciesLink. http://www.splink.org.br/index (last access in 20/10/2019)
http://www.splink.org.br/index... ). We considered only native taxa identified at the taxonomic level of species and excluded those with dubious identification, at the genera-level and exotic species (cited in a separate table - ^{Appendix 1} Supplementary Material The following online material is available for this article: Appendix 1 - - Excluded species from PFSP species list and the respective reason of exclusion. Appendix 2 - - Number of exclusive species among studies compared with this study. Appendix 3 - - Species occurring at 11 (50%) or more lists among the compared studies and PFSP. ). For angiosperms, the taxa classification followed the APG IV system (2016)APG IV. 2016. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV. Bot. J. Linn. Soc. 181(1):1-20., and for ferns or lycophytes, the PPG I system (2016)PPG I. 2016. A community‐derived classification for extant lycophytes and ferns. J. Syst. Evol. 54(6):563-603.. The nomenclature and synonymizations were updated according to the Flora do Brasil (2020)FLORA DO BRASIL. 2020. Jardim Botânico do Rio de Janeiro. http://floradobrasil.jbrj.gov.br/(last access in 12/05/2020)
http://floradobrasil.jbrj.gov.br/... . When the data from Flora do Brasil (2020)FLORA DO BRASIL. 2020. Jardim Botânico do Rio de Janeiro. http://floradobrasil.jbrj.gov.br/(last access in 12/05/2020)
http://floradobrasil.jbrj.gov.br/... were outdated or not available, we used the Tropicos (2019)TROPICOS. 2019. http://www.tropicos.org (last access in 25/10/2019)
http://www.tropicos.org... database.

The life forms of the species were obtained through field observations, assessing the herbarium specimen labels information and, when this was not possible, we consulted the Flora do Brasil (2020)FLORA DO BRASIL. 2020. Jardim Botânico do Rio de Janeiro. http://floradobrasil.jbrj.gov.br/(last access in 12/05/2020)
http://floradobrasil.jbrj.gov.br/... data. Shrubs and subshrubs were unified in a single category. To assess the number of exclusive and common species among the vegetation types in the study area, we built Venn diagrams for all the species together, and for each life form. The threatened species were classified according to The Red Book of Brazilian Flora (Martinelli & Moraes, 2013MARTINELLI, G. & MORAES, M.A. (eds). 2013. Livro Vermelho da Flora do Brasil. CNCFlora, Rio de Janeiro.) and the list of threatened flora of São Paulo (SMA-SP, 2016SMA-SP. 2016. Publica a segunda revisão da lista oficial das espécies da flora ameaçadas de extinção no estado de São Paulo. Resolução nº 57, de 5 de junho de 2016. Secretaria Estadual do Meio Ambiente, São Paulo.).

The similarity in species composition was estimated by the Jaccard Index (Mueller-Dombois & Ellenberg, 1974MUELLER-DOMBOIS, D. & ELLENBERG, H. 1974. Aims and methods of vegetation ecology. John Wiley & Sons, New York.), as well as this study and 14 others carried out in Cerrado sensu lato, SSF and RP vegetation types. Only vascular or phanerogamic floristic surveys were compared, which included studies performed in the South and Southeast regions of Brazil. Only native taxa identified at the taxonomic level of species were considered. Studies that sampled more than one vegetation type were subdivided into independent lists (e.g. Durigan et al., 1999DURIGAN, G., BACIC, M.C., FRANCO, G.A.D.C. & SIQUEIRA, M.F. 1999. Inventário florístico do cerrado na Estação Ecológica de Assis, SP. Hoehnea. 26(2):149-172.), resulting in 22 lists (Table 1). We used a binary matrix with presence (1) and absence (0) to conduct an agglomerative hierarchical clustering analysis (Legendre & Legendre, 2012LEGENDRE, P. & LEGENDRE, L. 2012. Numerical Ecology (Vol. 24). 3 ed. Elsevier, Amsterdam.), by employing UPGMA (average linkage), and Jaccard dissimilarity in the vegan package (Oksanen et al. 2013OKSANEN, J., BLANCHET, F.G., KINDT, R., LEGENDRE, P., MINCHIN, P.R., O’HARA, R.B., SIMPSON, G.L., SOLYMOS, P., STEVENS, M.H.H. & WAGNER, H. 2013. Vegan: Community Ecology Package. R Package Version 2.0-10. http://vegan.r-forge.r-project.org (last access in 28/10/2020)
http://vegan.r-forge.r-project.org... ) in R (R Core Team 2020R CORE TEAM. 2020. R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna. https://www.R-project.org/ (last access in 02/03/2020)
https://www.R-project.org/... ).

Results

A total of 684 species distributed in 107 families and 387 genera were recorded (640 angiosperms and 44 ferns or lycophytes) (Table 2). The richest families were Fabaceae, with 64 species, followed by Myrtaceae (41), Orchidaceae (39), Rubiaceae (37), Asteraceae (35), Bignoniaceae (26), Malvaceae (20), Euphorbiaceae (19), Malpighiaceae and Sapindaceae (17 each) (Figure 3). These families account for 46% of the surveyed species, and 35 families (32.7%) had one single species. The richest genera were Eugenia (19 species), Miconia (14), Myrcia (11), Solanum (nine), Peperomia and Serjania (eight each). A separate list shows 152 excluded species and the reason for the exclusion (^{Appendix 1} Supplementary Material The following online material is available for this article: Appendix 1 - - Excluded species from PFSP species list and the respective reason of exclusion. Appendix 2 - - Number of exclusive species among studies compared with this study. Appendix 3 - - Species occurring at 11 (50%) or more lists among the compared studies and PFSP. ). The richest vegetation type was SSF, with 478 species, followed by the CER (418) and the RP (231).

Figure 3
The most species-rich families in the Porto Ferreira State Park, Porto Ferreira, São Paulo state, southeastern Brazil.

From the registered species, 581 were listed in previous surveys, and 527 of them we found in herbaria vouchers. Added to the last floristic list prepared for the management plan, were 412 species, 70 of which were drawn from aleatory collections (not cited in previous surveys), and 33 from collections carried out by the authors of this study. Eight species recorded here are on the threatened list at the national level (Martinelli & Moraes, 2013MARTINELLI, G. & MORAES, M.A. (eds). 2013. Livro Vermelho da Flora do Brasil. CNCFlora, Rio de Janeiro.) two of which - Anemopaegma arvense (Vell.) Stellfeld ex de Souza and Cariniana legalis (Mart.) Kuntze - are endangered, and six species [Euterpe edulis Mart., Zeyheria tuberculosa (Vell.) Bureau ex Verl., Croton leptobotryus Müll.Arg., Cedrela fissilis Vell., Cattleya walkeriana Gardner and Isabelia virginalis Barb.Rodr.] are listed as vulnerable. The list of threatened São Paulo State flora (SMA 2016) follows the same species and threat category as on the national list, except for Cariniana legalis (Mart.) Kuntze, classified as vulnerable.The arboreal habit was the predominant life form in the PFSP (41.8%), followed by herbs (25.7%), shrubs/subshrubs (17.9%) and climbing plants (14.7%) (Figure 4). The results listed 84 species (12.2%) with two or more life forms. Concerning all life forms, 9.6%, 18.8% and 18.5% of the species are exclusive to RP, CER and SSF, respectively. The number of exclusive species for the CER is mainly related to the high number of trees species. However, herbs represented 40% and 50% of the exclusive species found in the SSF and RP, respectively.

Figure 4
Venn diagrams for vascular plants recorded in Porto Ferreira State Park, Porto Ferreira, São Paulo State, southeastern Brazil. RP: riparian forest, SSF: seasonal semidecidual forest; CER: cerradão.

The number of species compiled for the lists of the floristic similarity comparison was 2745, of which 1420 (51.7%) are exclusively from a single area (^{Appendix 2} Supplementary Material The following online material is available for this article: Appendix 1 - - Excluded species from PFSP species list and the respective reason of exclusion. Appendix 2 - - Number of exclusive species among studies compared with this study. Appendix 3 - - Species occurring at 11 (50%) or more lists among the compared studies and PFSP. ). Only 43 species (1.5%) occurred in 50% or more of the lists, and no species were recorded for all areas (^{Appendix 3} Supplementary Material The following online material is available for this article: Appendix 1 - - Excluded species from PFSP species list and the respective reason of exclusion. Appendix 2 - - Number of exclusive species among studies compared with this study. Appendix 3 - - Species occurring at 11 (50%) or more lists among the compared studies and PFSP. ). The dendrogram showed a high coefficient of cophenetic correlation of 0.82. The greater the distance was between areas, the greater the floristic dissimilarities among them. The cluster analysis showed the segregation of three major floristic groups (Figure 5). The first group (1) clustered an ecotone area and two RP (one of which belongs to the same survey as the ecotone); the second group (2) categorized the studies carried out in Cerrado vegetation types; and the third group (3) all vegetation types of the PFSP and areas under SSF formation.

Figure 5
Cluster analysis (UPGMA) using Jaccard index among fifteen studies carried out in Cerrado, SSF and riparian forest vegetation. Areas (identified by letters A-V) are given in Table 1. SSF: seasonal semidecidual forest; Cerrado s.l.: Cerrado sensu lato; Cerrado s.s.: Cerrado sensu stricto; RF-AF: riparian forest under Atlantic Forest ecoregion; RF-Cer: riparian forest under Cerrado ecoregion.

Discussion

The comprehensive checklist of the PFSP allowed us to show that the vascular flora in this ecotonal area is composed of a mixture of elements of both the Atlantic Forest and Cerrado biogeographical provinces. The soil features are a possible explanation for the different vegetation types, and variations on a small scale. The overall richness is impressive, considering the small size of park; however, the number of species of each vegetation type recorded is in accordance with previous studies in these vegetation types. An Atlantic Forest similarity was found for the PFSP flora as a whole. The same relationship was found considering the CER and SSF floras independently, demonstrating the strong influence of Atlantic Forest on the area. Perhaps the geographical location of the PFSP, which reaches both biogeographical provinces in the eastern region, could be a reasonable explanation for these results.

The species richness of the PFSP was higher than that found by Cielo-Filho et al. (2015)CIELO-FILHO, R., BAITELLO, J.B., ARZOLLA, F.A.R.D.P., VILELA, F.E.S.P., KANASHIRO, M.M., MATTOS, I.F.A., AGUIAR, O.T., SOUZA, S.C.P.M., PASTORE, J.A. , FRANCO, G.A.D.C. & ZIMBACK, L. 2015. A vegetação da Estação Ecológica de Avaré: subsídios para o plano de manejo. IF. Ser. Regist. 53:5-42. - with 265 species -, the only vascular flora surveyed in an ecotonal site among Cerrado and Atlantic Forest in southeastern Brazil. The overwhelming number of species recorded for the PFSP vascular flora is ranked fourth among studies conducted under seasonal formations, behind only those of Lombardi & Gonçalves (2000)LOMBARDI, J.A. & GONÇALVES, M. 2000. Composição florística de dois remanescentes de Mata Atlântica do sudeste de Minas Gerais, Brasil. Rev. Brasil. Bot. 23(3):255-282., Lombardi et al. (2012)LOMBARDI, J.A., CARVALHO, C.S., BIRAL, L., SAKA, M.N. & HIEDA, S.M. 2012. Vascular flora of Serra do Japi Biological Reserve, Jundiaí, southeastern Brazil. Rodriguésia. 63(2):333-340. and Forzza et al. (2014)FORZZA, R.C., PIFANO, D.S., OLIVEIRA-FILHO, A.T., MEIRELES, L.D., FARIA, P.L., SALIMENA, F.R., MYNSSEN, C.M. & PRADO, J. 2014. Flora vascular da Reserva Biológica da Represa do Grama, Minas Gerais, e sua relação florística com outras florestas do sudeste brasileiro. Rodriguésia. 65(2):275-292.. Species richness tends to peak in ecotonal areas (Kark & van Rensburg, 2006KARK S. & VAN RENSBURG B. J. 2006. Ecotones: marginal or central areas of transition? Isr. J. Ecol. Evol. 52:29-53.), likely because transitional areas hold species from two or more neighboring communities, and due to high spatial heterogeneity (Kark 2012KARK, S. 2012. Ecotones and Ecological Gradients. In Encyclopedia of Sustainability Science and Technology (R. A. Meyers, ed). Springer Science & Business Media, New York, p.147-160.). Furthermore, along with the increase in botanical collection in protected areas, there is an appeal to enhance the importance of these areas to protect threatened species (Colli-Silva et al., 2019COLLI-SILVA, M., IVANAUSKAS, N.M. & SOUZA, F.M. 2019. Diagnóstico do conhecimento da biodiversidade de plantas vasculares nas unidades de conservação do estado de São Paulo. Rodriguésia. 70:e04582017.). In this sense, we found eight threatened species at regional and national levels. Thus, the ecotonal condition of the PFSP between two biogeographical provinces may contribute to increase the species richness, also being home to important threatened species.

Studies have been emphasizing the need for the expansion of botanical collection in protected areas (Colli-Silva et al. 2016COLLI-SILVA, M., BEZERRA, T.L., FRANCO, G.A.D.C., IVANAUSKAS, N.M. & SOUZA, F.M. 2016. Registros de espécies vasculares em unidades de conservação e implicações para a lista da flora ameaçada de extinção no estado de São Paulo. Rodriguésia. 67(2):405-425., 2019COLLI-SILVA, M., IVANAUSKAS, N.M. & SOUZA, F.M. 2019. Diagnóstico do conhecimento da biodiversidade de plantas vasculares nas unidades de conservação do estado de São Paulo. Rodriguésia. 70:e04582017.). The addition of more than 65% of the species number since the last PFSP floristic list (São Paulo, 2003SÃO PAULO. 2003. Plano de manejo do Parque Estadual de Porto Ferreira, Instituto Florestal, São Paulo. https://www.infraestruturameioambiente.sp.gov.br/fundacaoflorestal/planos-de-manejo/planos-de-manejo-planos-concluidos/plano-de-manejo-pe-porto-ferreira/ (last access in 02/03/2020)
https://www.infraestruturameioambiente.s... ) is significant, showing the importance of continuous research, and fieldwork, to update the knowledge about the flora. Since the Colli-Silva et al. study of 2016, which recorded 292 species for the PFSP, 235 vouchers have been added to the PFSP, most likely due to the digitization of the collections, later available in digital platforms (CRIA, 2019CRIA. 2019. SpeciesLink. http://www.splink.org.br/index (last access in 20/10/2019)
http://www.splink.org.br/index... ), and to the addition of our own collections. Moreover, notably there are species that we found in the field, such as Cissus serroniana (Glaz.) Lombardi (Vitaceae), which had never before been recorded fertile, and, therefore, were not included in the checklist. Even though the PFSP has a significant number of studies concerning its flora, we predict that further collections will increase the number of species in this protected area.

Considering each vegetation type of the PFSP, the richness we found in the SSF averages roughly with that found in other studies (Corrêa et al., 2018CORRÊA, L.S., SCATIGNA, A.V., GISSI, D.S., SILVA, D.M., COTA, M.M.T., SOUZA, V.C., IVANAUSKAS, N.M., TAMASHIRO, J.Y. & RODRIGUES, R.R. 2018. Vascular flora checklist of the Ibicatu Ecological Station, Piracicaba, São Paulo, Brazil. Rev. Inst. Flor. 30(1):53-70.; Cielo-Filho et al., 2015CIELO-FILHO, R., BAITELLO, J.B., ARZOLLA, F.A.R.D.P., VILELA, F.E.S.P., KANASHIRO, M.M., MATTOS, I.F.A., AGUIAR, O.T., SOUZA, S.C.P.M., PASTORE, J.A. , FRANCO, G.A.D.C. & ZIMBACK, L. 2015. A vegetação da Estação Ecológica de Avaré: subsídios para o plano de manejo. IF. Ser. Regist. 53:5-42.; Forzza et al., 2014FORZZA, R.C., PIFANO, D.S., OLIVEIRA-FILHO, A.T., MEIRELES, L.D., FARIA, P.L., SALIMENA, F.R., MYNSSEN, C.M. & PRADO, J. 2014. Flora vascular da Reserva Biológica da Represa do Grama, Minas Gerais, e sua relação florística com outras florestas do sudeste brasileiro. Rodriguésia. 65(2):275-292.; Pifano et al., 2013PIFANO, D.S., GARCIA, P.O., VALENTE, A.S.M., ANTUNES, K. & CASTRO, R.M. 2013. Flora fanerogâmica da mata do sítio Primavera na APA Serra dos Núcleos, São João Nepomuceno, Minas Gerais e suas relações florísticas com florestas ombrófilas e semidecíduas no domínio atlântico. Biota. 6(3):12-39.; Rossetto & Vieira, 2013ROSSETTO, E.F.S. & VIEIRA, A.O.S. 2013. Vascular Flora of the Mata dos Godoy State Park, Londrina, Paraná, Brazil. Check List. 9(5):1020-1034.; Lombardi et al., 2012LOMBARDI, J.A., CARVALHO, C.S., BIRAL, L., SAKA, M.N. & HIEDA, S.M. 2012. Vascular flora of Serra do Japi Biological Reserve, Jundiaí, southeastern Brazil. Rodriguésia. 63(2):333-340.; Guaratini et al., 2008GUARATINI, M.T.G., GOMES, E.P.C., TAMASHIRO, J.Y. & RODRIGUES, R.R. 2008. Composição florística da Reserva Municipal de Santa Genebra, Campinas, SP. Rev. Brasil. Bot. 31(2):323-337.; Lombardi & Gonçalves, 2000LOMBARDI, J.A. & GONÇALVES, M. 2000. Composição florística de dois remanescentes de Mata Atlântica do sudeste de Minas Gerais, Brasil. Rev. Brasil. Bot. 23(3):255-282.). In the CER, the number of species recorded in our study is higher than other Cerrado surveys (Cavassan & Weiser, 2015CAVASSAN, O. & WEISER, V.L. 2015. Vascular flora of the cerrado of Bauru-SP. Biota Neotropica. 15(3): e20140093. http://dx.doi.org/10.1590/1676-0611-BN-2014-0093 (last access in 22/11/2020)
http://dx.doi.org/10.1590/1676-0611-BN-2... ; Cielo-Filho et al. 2015CIELO-FILHO, R., BAITELLO, J.B., ARZOLLA, F.A.R.D.P., VILELA, F.E.S.P., KANASHIRO, M.M., MATTOS, I.F.A., AGUIAR, O.T., SOUZA, S.C.P.M., PASTORE, J.A. , FRANCO, G.A.D.C. & ZIMBACK, L. 2015. A vegetação da Estação Ecológica de Avaré: subsídios para o plano de manejo. IF. Ser. Regist. 53:5-42.; Carvalho et al., 2010CARVALHO, M.B., ISHARA, K.L. & RODELLA, R.C.S.M. 2010. Vascular Flora of a Cerrado sensu stricto remnant in Pratânia, state of São Paulo, southeastern Brazil. Check List. 6(3):350-357.; Ishara et al., 2008; Batalha & Mantovani, 2001BATALHA, M.A. & MANTOVANI, W. 2001. Floristic composition of the cerrado in the Pé-de-gigante reserve (Santa Rita do Passa Quatro, Southeastern Brazil). Acta Bot. Bras. 15(3):289-304.; Durigan et al., 1999DURIGAN, G., BACIC, M.C., FRANCO, G.A.D.C. & SIQUEIRA, M.F. 1999. Inventário florístico do cerrado na Estação Ecológica de Assis, SP. Hoehnea. 26(2):149-172.; Batalha et al., 1997BATALHA, M.A., ARAGAKI, S. & MANTOVANI, W. 1997. Florística do cerrado em Emas (Pirassununga, SP). Bol. Bot. Univ. 16:49-64.). As for the RP, there are no studies of vascular flora carried out specifically on this vegetation type, but rather inevitably included as a part of a larger work, such as those of Durigan et al. (1999)DURIGAN, G., BACIC, M.C., FRANCO, G.A.D.C. & SIQUEIRA, M.F. 1999. Inventário florístico do cerrado na Estação Ecológica de Assis, SP. Hoehnea. 26(2):149-172. and Cielo-Filho et al. (2015)CIELO-FILHO, R., BAITELLO, J.B., ARZOLLA, F.A.R.D.P., VILELA, F.E.S.P., KANASHIRO, M.M., MATTOS, I.F.A., AGUIAR, O.T., SOUZA, S.C.P.M., PASTORE, J.A. , FRANCO, G.A.D.C. & ZIMBACK, L. 2015. A vegetação da Estação Ecológica de Avaré: subsídios para o plano de manejo. IF. Ser. Regist. 53:5-42., making comparison difficult.

The most species-rich families found in our study are also the richest ones for the Cerrado as a whole (Souza, et al. 2018SOUZA, V.C., FLORES, T.B., COLLETA, G.D. & COELHO, R.L.G. 2018. Guia das plantas do Cerrado. Editora Taxon Brasil, Piracicaba.), and seasonal forests of the Atlantic Forest (Souza, et al., 2019aSOUZA, V.C., TOLEDO, C.P., SAMPAIO, D., BÍGIO, N.C., COLLETA, G.D., IVANAUSKAS, N.M. & FLORES, T.B. 2019a. Guia das plantas da mata atlântica: floresta estacional. Liana Produções Editoriais, Piracicaba.). Orchidaceae, Fabaceae, Asteraceae and Myrtaceae are, respectively, the most diverse families in the Atlantic Forest (BFG, 2015BFG. 2015. Growing knowledge: an overview of Seed Plant diversity in Brazil. Rodriguésia. 66(4):1085-1113.). Thus, Fabaceae, Asteraceae and Orchidaceae constitute the three richest families in the Cerrado (Mendonça et al. 2008MENDONÇA, R.C., FELFILI, J.M., WALTER, B.M.T., SILVA JÚNIOR, M.C., REZENDE, A.V., FILGUEIRAS, T.S. & NOGUEIRA, P.E. 2008. Flora vascular do bioma Cerrado: Checklist com 12.356 espécies. In Cerrado: ecologia e flora - vol. 2 (S.M. Samo, S.P. Almeida & J.F. Ribeiro, eds). Embrapa, Planaltina, p.421-1279.). The richest genera found in the PFSP - Eugenia, Micona and Myrcia - are ranked as the genera with the highest number of tree species in the Cerrado and the Atlantic Forest, under semideciduous formation (Oliveira-Filho & Fontes 2000OLIVEIRA-FILHO, A.T. & FONTES, M.A.L. 2000. Patterns of Floristic Differentiation among Atlantic Rain Forests in Southeastern Brazil and the Influence of Climate. Biotropica. 32(4b):793-810.). Considering Brazilian flora as a whole, Eugenia, Miconia, Myrcia, Psychotria, Solanum and Peperomia are also classified among the 30 most diverse Angiosperm genera (BFG, 2015BFG. 2015. Growing knowledge: an overview of Seed Plant diversity in Brazil. Rodriguésia. 66(4):1085-1113.).

Generalist species, establishing and thriving both in the Cerrado and in the Atlantic Forest, generally become very abundant in transition areas between these two biogeographical provinces (Durigan et al. 2012DURIGAN, G., RAMOS, V.S., IVANAUSKAS, N.M. & FRANCO, G.A.D.C. 2012. Espécies indicadoras de fitofisionomias na transição cerrado-mata atlântica no estado de São Paulo. SMA/CBRN, São Paulo., Morrone, 2014MORRONE, J.J. 2014. Biogeographical regionalisation of the Neotropical region. Zootaxa. 3782:1-110., 2017MORRONE, J.J. 2017. Neotropical biogeography: regionalization and evolution. CRC Press, Boca Raton.). All species classified as generalists by Durigan et al. (2012)DURIGAN, G., RAMOS, V.S., IVANAUSKAS, N.M. & FRANCO, G.A.D.C. 2012. Espécies indicadoras de fitofisionomias na transição cerrado-mata atlântica no estado de São Paulo. SMA/CBRN, São Paulo. were found in the PFSP flora [e.g. Copaifera langsdorffii Desf., Cordia trichotoma (Vell.) Arráb. ex Steud, Mabea fistulifera Mart., Ocotea corymbosa (Meisn.) Mez, Platypodium elegans Vogel, Senegalia polyphyla (DC.) Britton & Rose, Terminalia glabrescens Mart. and Vochysia tucanorum Mart.]. Durigan et al. (2012)DURIGAN, G., RAMOS, V.S., IVANAUSKAS, N.M. & FRANCO, G.A.D.C. 2012. Espécies indicadoras de fitofisionomias na transição cerrado-mata atlântica no estado de São Paulo. SMA/CBRN, São Paulo. also pointed out 20 typical species of SSF, of which 15 were reported in the PFSP flora (e.g. Aspidosperma polyneuron Müll.Arg., Cedrela fissilis Vell., Metrodorea nigra A.St.-Hil. and Guarea kunthiana A. Juss.). In a study encompassing most of the Cerrado woody vegetation, Ratter et al. (2003)RATTER, J.A., BRIDGEWATER, S. & RIBEIRO, J.F. 2003. Analysis of the floristic composition of the Brazilian Cerrado vegetation. III: Comparision of the woody vegetation of 376 areas. Edinb. J. Bot. 60(1):57-109. found 38 species with the highest percentage of occurrence, of which 26 (68.4%) occur in PFSP CER (e.g. Qualea grandiflora Mart., Qualea parviflora Mart., Bowdichia virgilioides Kunth, Dimorphandra mollis Benth., Lafoensia pacari A.St.-Hil., Connarus suberosus Planch. and Hymenaea stigonocarpa Mart. ex Hayn). These results emphasize the PFSP as a species reservatory of several biogeographical entities, reinforcing its role in conservation.

The trees were the most species-rich life forms in the PFSP, following the patterns of forest vegetation types (IBGE, 2012IBGE. 2012. Manual técnico da vegetação brasileira. Segunda edição revista e ampliada. Série Manuais Técnicos em Geociências, Rio de Janeiro.; Forzza et al., 2014FORZZA, R.C., PIFANO, D.S., OLIVEIRA-FILHO, A.T., MEIRELES, L.D., FARIA, P.L., SALIMENA, F.R., MYNSSEN, C.M. & PRADO, J. 2014. Flora vascular da Reserva Biológica da Represa do Grama, Minas Gerais, e sua relação florística com outras florestas do sudeste brasileiro. Rodriguésia. 65(2):275-292.). Regarding the shared species among the vegetation types, trees perform a major contribution to the exclusive species of the CER. The different edaphic physicochemical features among the CER and SSF are the main causes of their differences, even in small scales (Gottsberger & Silberbauer-Gottsberger, 2006GOTTSBERGER, G. & SILBERBAUER-GOTTSBERGER, I. 2006. Life in the Cerrado: a South American Tropical Seasonal Vegetation, Vol. II. Pollination and seed dispersal. Reta Verlag, Ulm.; Pinheiro et al. 2009PINHEIRO, M.H.O., ARANTES, S.A.C.M., JIMENEZ-RUED, J.R. & MONTEIRO, R. 2009. Caracterização edáfica de um ecótono savânico-florestal no sudeste brasileiro. Iheringia Ser. Bot. 64(2):15-24.). Edaphic factors, such as water availability in the soil, and soil composition, may play a role in vegetation type differentiation (Haridasan 1992HARIDASAN, M. 1992. Observations on soils, foliar nutrient concentration and floristic composition of cerrado sensu stricto and cerradão communities in central Brazil. In Nature and Dynamics of Forest: savanna boundaries (P.A. Furley, J. Proctor & J.A. Ratter, eds). Chapman & Hall, London, p.171-184.). The heterogeneity of the soil, on a small-scale, can also act as a plant community ecological driver, increasing diversity and promoting structural changes (Souza et al., 2019bSOUZA, C.R., MOREL, J.D., SANTOS, A.B.M., DA SILVA, W.B., MAIA, V.A., COELHO, P.A., REZENDE, V.L. & DOS SANTOS, R.M. 2019b. Small-scale edaphic heterogeneity as a floristic-structural complexity driver in Seasonally Dry Tropical Forests tree communities. J. For. Res. 31:2347-2357.). In a study carried out on the PFSP soil and its relationship to the vegetation, Rossi et al. (2005)ROSSI, M., MATTOS, I.F.A., COELHO, R.M., MENK, J.R.F., ROCHA, F.T., PFEIFER, R.M. & DEMARIA, I.C. 2005. Relação solos/vegetação em área natural no Parque Estadual de Porto Ferreira, São Paulo. Rev. Inst. Flor. 17(1):45-61. found several types of soil. The authors also found an association between CER, SSF and RP vegetation and dystrophic oxisols, eutrophic argisols and dystrophic fluvial neosols, respectively.

A substantial number of herbs species were found in the PFSP, including the epiphytic and terricolous habit. The impressive importance of the herbs in the PFSP is most likely due to the contribution of the Cerrado elements; usually the number recorded in the AF is lower than the one we recorded here (Vieira et al., 2015VIEIRA, L.T.A., POLISEL R.T., IVANAUSKAS, N.M., SHEPHERD, G.J., WAECHTER, J.L., YAMAMOTO, K. & MARTINS, F.R. 2015. Geographical patterns of terrestrial herbs: a new component in planning the conservation of the Brazilian Atlantic Forest. Biodivers. Conserv. 24(9):2181-2198.). An explanation for this may be the ecotonal feature of the PFSP, and the high number of shared species between the CER and SSF. Thus, even in the Cerrado forestry vegetation type, the herbaceous layers must be considered as an important synusia. On the other hand, the high number of exclusive herbaceous species in the RP must be attributed to the microclimatic conditions propitious to the epiphytes’ development (Marcusso et al., 2016MARCUSSO, G.M., DICKFELDT, E.P., BERTONI, J.E.A. & MONTEIRO, R. 2016. Epífitas vasculares do Parque Estadual de Porto Ferreira, São Paulo, Brasil. Rev. Inst. Flor. 28(2):119-133.). The number of species of climbing plants in the present study was lower than that found by Vargas et al. (2018)VARGAS, B.C., OLIVEIRA, A.P.C., UDULUTSCH, R.G., MARCUSSO, G. M., SABINO, G.P., MELO, P.H.A., GRILLO, R.M.M., KAMIMURA, V.A. & ASSIS, M.A. 2018. Climbing plants of Porto Ferreira State Park, southeastern Brazil. Biota Neotropica. 18(2):e20170346. https://doi.org/10.1590/1676-0611-bn-2017-0346 (last access in 10/02/2020)
https://doi.org/10.1590/1676-0611-bn-201... , because we considered only taxa identified at the taxonomic level of species and, occasionally, used the life form classification according to the Flora do Brasil (2020)FLORA DO BRASIL. 2020. Jardim Botânico do Rio de Janeiro. http://floradobrasil.jbrj.gov.br/(last access in 12/05/2020)
http://floradobrasil.jbrj.gov.br/... . This classification does not always reflect Vargas’ et al. (2018)VARGAS, B.C., OLIVEIRA, A.P.C., UDULUTSCH, R.G., MARCUSSO, G. M., SABINO, G.P., MELO, P.H.A., GRILLO, R.M.M., KAMIMURA, V.A. & ASSIS, M.A. 2018. Climbing plants of Porto Ferreira State Park, southeastern Brazil. Biota Neotropica. 18(2):e20170346. https://doi.org/10.1590/1676-0611-bn-2017-0346 (last access in 10/02/2020)
https://doi.org/10.1590/1676-0611-bn-201... interpretations.

We found an outstanding heterogeneity among the floristic surveys analyzed, in which very few species occurred in 50% or more of the areas, and no species occurred in all areas. This may be due to the addition of surveys from different vegetation types belonging to the Cerrado and the Atlantic Forest. Floristic comparisons are usually made between vegetation types within the same biogeographical provinces (e.g. Carvalho et al. 2010CARVALHO, M.B., ISHARA, K.L. & RODELLA, R.C.S.M. 2010. Vascular Flora of a Cerrado sensu stricto remnant in Pratânia, state of São Paulo, southeastern Brazil. Check List. 6(3):350-357.; Ishara et al., 2008; Guaratini et al., 2008GUARATINI, M.T.G., GOMES, E.P.C., TAMASHIRO, J.Y. & RODRIGUES, R.R. 2008. Composição florística da Reserva Municipal de Santa Genebra, Campinas, SP. Rev. Brasil. Bot. 31(2):323-337.; Pifano et al., 2013PIFANO, D.S., GARCIA, P.O., VALENTE, A.S.M., ANTUNES, K. & CASTRO, R.M. 2013. Flora fanerogâmica da mata do sítio Primavera na APA Serra dos Núcleos, São João Nepomuceno, Minas Gerais e suas relações florísticas com florestas ombrófilas e semidecíduas no domínio atlântico. Biota. 6(3):12-39.). Here, including areas of both Cerrado and Atlantic Forest, we found that the areas with the same vegetation type are more inclined to be floristically similar. However, in ecotones, areas with different vegetation types in the same location tend to share more species with each other than areas in which the same vegetation type occurs at a distance from one another. These results have been reported for several studies (e.g. Oliveira-Filho & Fontes, 2000OLIVEIRA-FILHO, A.T. & FONTES, M.A.L. 2000. Patterns of Floristic Differentiation among Atlantic Rain Forests in Southeastern Brazil and the Influence of Climate. Biotropica. 32(4b):793-810.; Oliveira-Filho et al., 2015OLIVEIRA-FILHO, A.T., BUDKE, J.C., JARENKOW, J.A., EISENLOHR, P.V. & NEVES, D.R.M. 2015. Delving into the variations in tree species composition and richness across South American subtropical Atlantic and Pampean forests. J. Plant. Ecol. 8(3):242-226.; Dryflor et al., 2016DRYFLOR, B.R.K., DELGADO-SALINAS, A., DEXTER, K.G., LINARES-PALOMINO, R., OLIVEIRA-FILHO, A., PRADO, D. et al. 2016. Plant diversity patterns in neotropical dry forests and their conservation implications. Science. 353(6306):1383-1387.), contrary to the general biogeographic hypothesis that similar species would be shared between regions due to the restriction of their dispersion capacity (Lieberman, 2003LIEBERMAN, B.S. 2003. Unifying Theory and Methodology in Biogeography. In Evolutionary Biology (R.J. MacIntyre & M.T. Clegg, eds). Kluwer Academic/Plenum Publishers, New York, p.1-25.). Therefore, the fact that the CER of the PFSP joined in the SSF group might be due to the high spatial proximity correlation between them, suggesting a strong relationship with this matrix (Pinheiro & Monteiro, 2008PINHEIRO, M.H.O. & MONTEIRO, R. 2008. Florística de uma Floresta Estacional Semidecidual, localizada em ecótono savânico-florestal, no município de Bauru, SP, Brasil. Acta Bot. Bras. 22(4):1085-1094.; Passos et al., 2018PASSOS, F.B., MARIMON, B.S., PHILLIPS, O.L., MORANDI, P.S., NEVES, E.C., ELIAS, F., REIS, S.M., OLIVEIRA, B., FELDPAUSCH, T.R. & MARIMON JÚNIOR, B.H. 2018. Savanna turning into forest: concerted vegetation change at the ecotone between the Amazon and ‘‘Cerrado’’ biomes. Braz. J. Bot. 41(3):611-619.), with the SSF exerting more influence over the CER flora.

The PFSP harbours eight threatened species, and a huge number of vascular plant species. We also found a greater floristic similarity within the PFSP, and studies carried out in the Atlantic Forest. There is a lack of floristic studies that include all life forms conducted in ecotonal areas in southeastern Brazil. The reason for this, we believe, is due to the difficulty in recognizing an ecotone, and the negligence on supporting descriptive studies. This is an unprecedented floristic list recorded for the PFSP. The overwhelming diversity and the pool of species found, highlights this remnant as an important protected area. Results shown here can be used as a floristic reference for future conservation measures, and restorative actions in areas under ecotonal influences of the Atlantic Forest and the Cerrado.

Acknowledgments

We would like to thank Reinaldo Monteiro and André Luis Teixeira de Lucca for contributing with the flora surveys efforts in PFSP; Jomar Jardim (UFSB), Otávio Luis Marques da Silva (IBt-SP) and Júlio Antonio Lombardi (UNESP) for helping us with some identifications. We are grateful to PFSP staff for supporting fieldwork.

Supplementary Material

The following online material is available for this article:

Appendix 1 - - Excluded species from PFSP species list and the respective reason of exclusion.

Appendix 2 - - Number of exclusive species among studies compared with this study.

Appendix 3 - - Species occurring at 11 (50%) or more lists among the compared studies and PFSP.

References

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