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Flora of Espírito Santo: Oleandraceae

Abstract

As part of the Flora of Espírito Santo project, we present the taxonomic treatment of Oleandraceae, with key, descriptions, illustrations, and list of specimens examined. In this state, two species of Oleandra occur: O. brasiliana and O. quartziticola. Apparently, O. quartziticola is narrow endemic, whereas O. brasiliana is widespread in the Brazilian Atlantic Rainforest. Both species occur in the Southern part of the state.

Key words:
endemic ferns; epipetric ferns; Oleandra; Southeastern Brazil

Resumo

Como parte do projeto Flora do Espírito Santo, aqui é apresentado o tratamento taxonômico de Oleandraceae, contendo chave de identificação, descrições, ilustrações e lista de espécimes examinados. Neste estado, duas espécies de Oleandra ocorrem: O. brasiliana e O. quartziticola. Aparentemente, O. quartziticola é micro-endêmica, enquanto O. brasiliana é amplamente distribuída na Floresta Atlântica. Ambas espécies ocorrem na porção sul do estado.

Palavras-chave:
samambaias endêmicas; samambaias epipétricas; Oleandra; Sudeste do Brasil

Introduction

Oleandraceae is a pantropical, monogeneric family of ferns with ca. 30 species currently recognized (Maxon 1914Maxon WR (1914) The American species of Oleandra. Contributions from the United States National Herbarium 17: 392-398.; Roux 2009Roux JP (2009) Synopsis of the Lycopodiophyta and Pteridophyta of Africa, Madagascar and neighbouring islands. Strelitzia 23: 1-293.; Hovenkamp & Ho 2012Hovenkamp P & Ho B-C (2012) A revision of the fern genus Oleandra (Oleandraceae) in Asia. Phytokeys 11: 1-37.; Zhang & Hovenkamp 2013Zhang X & Hovenkamp P (2013) Oleandraceae. In: Wu Z, Raven PH & Hong D (eds.) Flora of China. Vols. 2,3 (Lycopodiaceae through Polypodiaceae). Science Press, Beijing, Missouri Botanical Garden Press, St. Louis. Pp. 747-748.; Schwartsburd et al. 2016Schwartsburd PB, Miranda CV & Prado J (2016) Oleandra (Oleandraceae) in the Brazilian Atlantic Forest. American Fern Journal 106: 191-205.). This family is sister group to a clade containing Polypodiaceae and Davalliaceae. These three families essentially present epiphytic or epipetric habit, and abscission of leaves due to articulation on phyllopodia (Schneider et al. 2004Schneider H, Smith AR, Cranfill R, Hildebrand TJ, Haufler CH & Ranker TA (2004) Unraveling the phylogeny of polygrammoid ferns (Polypodiaceae and Grammitidaceae): exploring aspects of the diversification of epiphytic plants. Molecular Phylogenetics and Evolution 31: 1041-1063.; Tsutsumi & Kato 2006Tsutsumi C & Kato M (2006) Evolution of epiphytes in Davalliaceae and related ferns. Botanical Journal of the Linnean Society 151: 495-510.; Schuettpelz & Pryer 2008Schuettpelz E & Pryer KM (2008) Fern phylogeny. In: Ranker TA & Haufler CH (eds.) Biology and evolution of ferns and lycophytes. Cambridge University Press, Cambridge. Pp. 395-416.; Smith et al. 2008Smith AR, Pryer KM, Schuettpelz E, Korall P, Schneider H & Wolf PG (2008) Fern classification. In: Ranker TA & Haufler CH (eds.) Biology and evolution of ferns and lycophytes. Cambridge University Press, Cambridge. Pp. 417-467.). However, Oleandraceae is characterized by stipe-like phyllopodia and reniform indusia (Hovenkamp & Ho 2012Hovenkamp P & Ho B-C (2012) A revision of the fern genus Oleandra (Oleandraceae) in Asia. Phytokeys 11: 1-37.), whereas the phyllopodia in the other two families are rhizomatous (mostly lost in the Grammitids), and the indusia in Davalliaceae is generally cup-shaped and absent in Polypodiaceae (Smith et al. 2008Smith AR, Pryer KM, Schuettpelz E, Korall P, Schneider H & Wolf PG (2008) Fern classification. In: Ranker TA & Haufler CH (eds.) Biology and evolution of ferns and lycophytes. Cambridge University Press, Cambridge. Pp. 417-467.).

The number of species in Oleandra has been controversial, especially due to two conflicting species concept of Maxon (1914)Maxon WR (1914) The American species of Oleandra. Contributions from the United States National Herbarium 17: 392-398. and Tryon (1997Tryon RM (1997) Systematic notes on Oleandra. Rhodora 99: 335-343., 2000Tryon RM (2000) Systematic notes on the Old World fern genus Oleandra. Rhodora 102: 428-438.). In the Brazilian Atlantic Rainforest, one name has been largely adopted, Oleandra articulata (Sw.) C.Presl, but other two names have also been used, O. baetaeDamazio and O. hirta Brack. (e.g., Damazio 1906Damazio L (1906) Une nouvelle fougère du Brésil. Bulletin de l'Herbier Boissier, ser. 2, 6: 829.; Maxon 1914Maxon WR (1914) The American species of Oleandra. Contributions from the United States National Herbarium 17: 392-398.; Lisbôa 1956Lisbôa MA (1956) Ptéridophytes de Ouro Preto. Anais da Escola de Minas de Ouro Preto 29: 21-77.; Sehnem 1979Sehnem A (1979) Davaliáceas. In: Reitz R (ed.) Flora Ilustrada Catarinense. Herbário Barbosa Rodrigues, Itajaí, part I, fascicle DAVA. Pp. 1-18.; Salino & Almeida 2009Salino A & Almeida TE (2009) Oleandraceae. In: Stehmann JR et al. (eds.) Plantas da Floresta Atlântica. Instituto de Pesquisas Jardim Botânico do Rio de Janeiro, Rio de Janeiro. Pp. 98.; Sylvestre 2010Sylvestre LS (2010) Oleandraceae. In: Forzza RC et al. (orgs.) Catálogo das plantas e fungos do Brasil. Vol. 1. Instituto de Pesquisas Jardim Botânico do Rio de Janeiro, Rio de Janeiro. Pp. 546.; 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 CGV, 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.). Recently, Schwartsburd et al. (2016)Schwartsburd PB, Miranda CV & Prado J (2016) Oleandra (Oleandraceae) in the Brazilian Atlantic Forest. American Fern Journal 106: 191-205. proposed a narrower species concept for the Atlantic Rainforest species, considering five species and a putative hybrid there. These authors also indicated that the largely used name, O. articulata, do not apply to the species from the Atlantic Rainforest. For the state of Espírito Santo, they listed only O. brasiliana Schwartsb. & J.Prado and O. quartziticola Schwartsb. & J.Prado. For the project Flora of Espírito Santo, we here present the taxonomic treatment of Oleandraceae.

Material and Methods

This paper is based on the recently published taxonomic revision of Oleandra from the Brazilian Atlantic Rainforest, in which the following herbaria were visited: CESJ, MBM, OUPR, RB, SP, SPF, UPCB, and VIC, plus images online from K, NY, and US. In addition to these herbaria, we further visited MBML and VIES.

We studied Oleandra quartziticola in the field, in the municipality of Vargem Alta. Other species (including O. brasiliana) were also studied in the field, but not in Espírito Santo (see Schawrtsburd et al. 2016).

The distribution map was drawn using the software Diva-GIS (Hijmans 2015Hijmans RJ (2015) DIVA-GIS, a geographic information system for the analysis of biodiversity data. Version 7.5. Available at <http://www.diva.gis.org>. Access on 17 november 2016.
http://www.diva.gis.org...
). For the materials without geographical coordinates, we estimated them using Google Earth (<https://www.google.com/earth/>).

Results and Discussion

In the state of Espírito Santo, two species of Oleandra occur: O. brasiliana and O. quartziticola, and both are known only from a few localities in the southern part of the state (Fig. 1), and from few herbaria materials. Apparently, O. quartziticola is narrow endemic, whereas O. brasiliana is widespread in the Brazilian Atlantic Rainforest.

Figure 1
Distribution of Oleandra species in Espírito Santo and near areas - O. brasiliana (●); O. quartziticola (○). Altitudinal layer in gray scale for each 500 m, from white to black.

Oleandraceae

Plants epiphytic or rupestrial, rarely terrestrial. Rhizomes long-creeping or scrambling, with or without pruinose exsudates, 1-8 mm diam., with or without rhizophore-like roots, scaly; scales spreading or appressed, peltate, commonly glandular-ciliate. Phyllopodia stipe-like. Fronds sparse or in whorls, monomorphic, articulated to phyllopodia; petioles terete, glabrous, with scales and/or hairs; laminae simple, linear, elliptic or obovate; midribs glabrous, glabrescent, with scales, catenate hairs, and/or with glandular hairs; veins free or rarely partly anastomosing, with or without catenate and/or glandular hairs; laminar tissue between the veins with or without catenate and/or glandular hairs; laminar margins with or without catenate and/or glandular hairs; sori abaxial, in one row between midribs and laminar margins; indusia reniform or orbicular-reniform, glabrous or with hairs; spores monolete.

Oleandra Cav., Anales Hist. Nat. 1(2):115. 1799.

A pantropical genus with ca. 30 species, and no apparent center of diversity (ca. eight in Asia, four in Africa, and 15 in the Americas) (Maxon 1914Maxon WR (1914) The American species of Oleandra. Contributions from the United States National Herbarium 17: 392-398.; Roux 2009Roux JP (2009) Synopsis of the Lycopodiophyta and Pteridophyta of Africa, Madagascar and neighbouring islands. Strelitzia 23: 1-293.; Hovenkamp & Ho 2012Hovenkamp P & Ho B-C (2012) A revision of the fern genus Oleandra (Oleandraceae) in Asia. Phytokeys 11: 1-37.; Schwartsburd et al. 2016Schwartsburd PB, Miranda CV & Prado J (2016) Oleandra (Oleandraceae) in the Brazilian Atlantic Forest. American Fern Journal 106: 191-205.). In the state of Espírito Santo, two species occur as rupestrial in the mountains of quartz of the Southern part (Fig. 2a-j).

Figure 2
a-d. Oleandra quartziticola - a. habit; b. rhizome scale; c. detail of lamina abaxially and laminar margin; d. detail of sori and indusium. e-j. O. brasiliana - e. habit; f. habit; g. rhizome scale; h. articulation between phyllopodium and petiole; i. detail of lamina abaxially and laminar margin (from a mature glabrescent specimen); j. detail of sori and indusium. (a-d. Schwartsburd 3522; e,h-j. Schwartsburd 2516; f. Duarte 9767; g. Matos 377).

    Key to species of Oleandra from the state of Espírito Santo
  • 1. Laminae glabrescent (with scattered hairs on midribs and laminar margins, especially in the laminar apex), or appearing glabrous; indusia monochromatic.......................................1. Oleandra brasiliana

  • 1'. Laminae pubescent throughout (with conspicuous hairs on midribs, veins, and laminar margins); indusia bicolorous........................................................................................................2. Oleandra quartziticola

1. Oleandra brasiliana Schwartsb. & J.Prado, Amer. Fern J. 106(3): 199-201, f. 1e-j, 3. 2016. Figs. 1; 2e-j

Plants rupestrial or rarely epiphytic. Rhizomes light brown to blackish, not pruinose, 1.5-2.5 mm diam., with rhizophore-like roots; scales spreading, bicolorous, centrally black, orange at the margins, moderate to densely glandular-ciliate. Phyllopodia 10-40 mm long. Fronds sparse, 12-35 cm long; petioles 3-7(-10) cm long, shining dark brown or greenish-brown; laminae generally obovate, rarely elliptic, cuneate at base, aristate at apex, 10-30 × (2-)4.5-7 cm; midribs glabrescent on both sides, with scattered 2-5-celled catenate hairs and 2-5-celled glandular hairs, sometimes seeming glabrous, lacking scales; veins free, glabrous on both sides; laminar tissue between the veins abaxially glabrous, adaxially glabrescent, with scattered catenate and glandular hairs; laminar margins cartilagineous, glabrescent, with scattered catenate and glandular hairs, especially common at the laminar apices; sori infra-medial to medial; indusia orbicular-reniform, dark brown, glabrous; spores light brown.

Specimens examined: Cachoeiro de Itapemirim, Vargem Alta, Morro de Sal, 21.VIII.1948, A.C. Brade 19324 (RB-on 2 sheets); 21.VIII.1948, A.C. Brade 3452 (CESJ); Vargem Alta, 3.V.1966, A.P. Duarte 9767 (MBM, RB-on 2 sheets, VIC).

Additional specimens examined: BRAZIL. BAHIA: Arataca, Serra do Peito de Moça, RPPN, Caminho das Pedras, 15o10'25"S, 39o20'30"W, 950 m, 6.VIII.2006, P.H. Labiak et al. 3652 (NY, SP, UPCB). CEARÁ: Maranguape, 3o54'5"S, 38o43'12"W, 900 m, 10.IV.2011, P.B. Schwartsburd & J.A.P. Araújo 2516 (EAC, NY, SP, VIC). MINAS GERAIS: Santa Bárbara do Monte Verde, 21o57'55"S, 43o49'51"W, 1,200 m, 24.IV.2004, J.P.S. Condack 135 (RB, VIC). RIO DE JANEIRO: Serra do Couto, A. Glaziou 2153 (RB-30604 [on 2 sheets]). SÃO PAULO: Serra de Paranapiacaba, IX.1925, A.C. Brade 8402 (MBM, RB-on 2 sheets).

Oleandra brasiliana is widespread along the Brazilian Atlantic Rainforest, from the state of Bahia to São Paulo, with a disjunct population in Ceará. In the state of Espírito Santo it has been found only in the mountains of quartz (morros de sal) of Vargem Alta (previously belonging to Municipality of Cachoeiro do Itapemirim), as rupestrial on quartzitic boulders. Outside Espírito Santo, O. brasiliana also occurs as epiphyte and rupestrial in other types of rocks.

Oleandra brasiliana have commonly been identified as O. articulata.Schwartsburd et al. (2016)Schwartsburd PB, Miranda CV & Prado J (2016) Oleandra (Oleandraceae) in the Brazilian Atlantic Forest. American Fern Journal 106: 191-205. recently showed that O. articulata do not occur in the Brazilian Atlantic Rainforest.

Oleandra brasiliana (Fig. 2e-j) is characterized by its glabrescent laminae, sometimes appearing fully glabrous, generally obovate laminae, and lack of scales on midribs. It differs from O. quartziticola (Fig. 2a-d) by glabrescent laminae (vs. pubescent throughout), generally obovate laminae (vs. elliptical), glabrous veins (vs. pubescent on both sides), and monochromatic, dark brown indusia (vs. indusia bicolorous, centrally dark brown, light brown at the margins).

2. Oleandra quartziticola Schwartsb. & J.Prado, Amer. Fern J. 106(3): 202-203, f. 2g-j, 3. 2016. Figs. 1; 2a-d

Plants rupestrial. Rhizomes light brown, not pruinose, 1-3.5 mm diam. with rhizophore-like roots; scales spreading, bicolorous, centrally black, orange at the margins, densely glandular-ciliate. Phyllopodia 15-20 mm long. Fronds sparse, 18-30 cm long; petioles (2-)3-5.5 cm long, stramineous to brown; laminae elliptic, cuneate at base, acuminate or aristate at apex, 15-25 × 2-4.2 cm; midribs with 4-7-celled catenate hairs and 4-7-celled glandular hairs on both sides, lacking scales; veins free, with catenate and glandular hairs on both sides; laminar tissue between the veins abaxially glabrous, adaxially with sparse catenate and glandular hairs; laminar margins cartilagineous, with glandular hairs and a few catenate hairs; sori infra-medial; indusia orbicular-reniform, bicolorous, centrally dark brown, light brown at the margins, glabrous; spores light brown.

Specimens examined: Vargem Alta, ES-164, Km 332, 20o39'26.7"S, 41o0'20"W, 663 m, 23.I.2011, J.R. Pirani et al. 6209 (SP, SPF); Morro Branco, 20o39'28"S, 41o0'18"W, 640 m, 17.IX.2015, P.B. Schwartsburd & C.V. Miranda 3522 (FI, NY, RB, SP, UC, UPCB, VIC-on 2 sheets); 20o39'26"S, 41o0'19"W, 660 m, 17.IX.2015, P.B. Schwartsburd & C.V. Miranda 3525 (SP, VIC).

Oleandra quartziticola is known only from three populations at the type locality. Apparently, it is a narrow endemic species, but further collections are needed, especially in Southern Espírito Santo. It has been found in the same habitat of O. brasiliana: in the mountains of quartz (morros de sal) of Vargem Alta, as rupestrial on quartzitic boulders.

  • Editora de área: Dra. Valquíria Dutra

Acknowledgements

We thank the Curators and staff of herbaria MBML and VIES, Cecília V. Miranda (VIC) for helping in the field trip, Reinaldo Pinto for the illustrations, Fábio A.R. Matos for helping with figure compositions, and Robbin C. Moran (NY) for sending images of exsiccates from NY.

References

  • Damazio L (1906) Une nouvelle fougère du Brésil. Bulletin de l'Herbier Boissier, ser. 2, 6: 829.
  • Hijmans RJ (2015) DIVA-GIS, a geographic information system for the analysis of biodiversity data. Version 7.5. Available at <http://www.diva.gis.org>. Access on 17 november 2016.
    » http://www.diva.gis.org
  • Hovenkamp P & Ho B-C (2012) A revision of the fern genus Oleandra (Oleandraceae) in Asia. Phytokeys 11: 1-37.
  • Lisbôa MA (1956) Ptéridophytes de Ouro Preto. Anais da Escola de Minas de Ouro Preto 29: 21-77.
  • Maxon WR (1914) The American species of Oleandra Contributions from the United States National Herbarium 17: 392-398.
  • 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 CGV, 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.
  • Roux JP (2009) Synopsis of the Lycopodiophyta and Pteridophyta of Africa, Madagascar and neighbouring islands. Strelitzia 23: 1-293.
  • Salino A & Almeida TE (2009) Oleandraceae. In: Stehmann JR et al (eds.) Plantas da Floresta Atlântica. Instituto de Pesquisas Jardim Botânico do Rio de Janeiro, Rio de Janeiro. Pp. 98.
  • Schneider H, Smith AR, Cranfill R, Hildebrand TJ, Haufler CH & Ranker TA (2004) Unraveling the phylogeny of polygrammoid ferns (Polypodiaceae and Grammitidaceae): exploring aspects of the diversification of epiphytic plants. Molecular Phylogenetics and Evolution 31: 1041-1063.
  • Schuettpelz E & Pryer KM (2008) Fern phylogeny. In: Ranker TA & Haufler CH (eds.) Biology and evolution of ferns and lycophytes. Cambridge University Press, Cambridge. Pp. 395-416.
  • Schwartsburd PB, Miranda CV & Prado J (2016) Oleandra (Oleandraceae) in the Brazilian Atlantic Forest. American Fern Journal 106: 191-205.
  • Sehnem A (1979) Davaliáceas. In: Reitz R (ed.) Flora Ilustrada Catarinense. Herbário Barbosa Rodrigues, Itajaí, part I, fascicle DAVA. Pp. 1-18.
  • Smith AR, Pryer KM, Schuettpelz E, Korall P, Schneider H & Wolf PG (2008) Fern classification. In: Ranker TA & Haufler CH (eds.) Biology and evolution of ferns and lycophytes. Cambridge University Press, Cambridge. Pp. 417-467.
  • Sylvestre LS (2010) Oleandraceae. In: Forzza RC et al (orgs.) Catálogo das plantas e fungos do Brasil. Vol. 1. Instituto de Pesquisas Jardim Botânico do Rio de Janeiro, Rio de Janeiro. Pp. 546.
  • Tryon RM (1997) Systematic notes on Oleandra Rhodora 99: 335-343.
  • Tryon RM (2000) Systematic notes on the Old World fern genus Oleandra Rhodora 102: 428-438.
  • Tsutsumi C & Kato M (2006) Evolution of epiphytes in Davalliaceae and related ferns. Botanical Journal of the Linnean Society 151: 495-510.
  • Zhang X & Hovenkamp P (2013) Oleandraceae. In: Wu Z, Raven PH & Hong D (eds.) Flora of China. Vols. 2,3 (Lycopodiaceae through Polypodiaceae). Science Press, Beijing, Missouri Botanical Garden Press, St. Louis. Pp. 747-748.

Publication Dates

  • Publication in this collection
    Oct-Dec 2017

History

  • Received
    18 Nov 2016
  • Accepted
    03 Apr 2017
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