Spores of Serpocaulon (Polypodiaceae): morphometric and phylogenetic analyses

Abstract

The morphometry and sculpture pattern of Serpocaulon spores was studied in a phylogenetic context. The species studied were those used in a published phylogenetic analysis based on chloroplast DNA regions. Four additional Polypodiaceae species were examined for comparative purposes. We used scanning electron microscopy to image 580 specimens of spores from 29 species of the 48 recognised taxa. Four discrete and ten continuous characters were scored for each species and optimised on to the previously published molecular tree. Canonical correspondence analysis (CCA) showed that verrucae width/verrucae length and verrucae width/spore length index and outline were the most important morphological characters. The first two axes explain, respectively, 56.3% and 20.5% of the total variance. Regular depressed and irregular prominent verrucae were present in derived species. However, the morphology does not support any molecular clades. According to our analyses, the evolutionary pathway of the ornamentation of the spores is represented by depressed irregularly verrucae to folded perispore to depressed regular verrucae to irregularly prominent verrucae.

Keywords:

• character evolution
• ferns
• eupolypods I
• canonical correspondence analysis

Acknowledgements

The authors especially thank Carlos Jaramillo (Smithsonian Tropical Research Institute [STRI], Panamá) and Andres Pardo Trujillo (IIES, Colombia) for their kind support. The authors would like to thank colleagues at the COL, CAUP, HUA, FAUC, PSO and UPCB herbaria for allowing to study the collections, and to Jorge Ceballos (STRI) for his assistance in the SEM laboratory, Maria José Leaño (UNIANDES) for his graphic assistance and Simon Quintero (UNIANDES) for special support. Sincere thanks to Michael Sundue (UVM), Lucas Majure (DBG), Robbin Moran (NYBG) and the anonymous reviewers for providing constructive comments. This work was supported by the Centre of Paleoecology and Archaeology (CTPA) at the STRI and Caldas University (IIES Institute and FAUC Herbarium).

Disclosure statement

No potential conflict of interest was reported by the authors.

Specimens examined

Serpocaulon appressum (Copel.) A. R. Sm., Bolivia, Provincia de Nor Yungas: Municipio de Coroico, Parque Nacional Cota Pata, Camino antiguo de Sillutinkara, 2444 m, 16° 15ʹ S, 67° 53ʹ W, 15 January 2012, Sanín et al. 5535 (HUA).

Serpocaulon attenuatum (C. Presl) A. R. Sm., Colombia, Meta: Reserva Natural la Macarena, margen izquierdo del río Guayabero, 1er raudal, 7 June 1970, Echeverry 2037 (COL).

Serpocaulon caceresii (Sodiro) A. R. Sm., Colombia, Antioquia, Cocorná, vereda La Piñuela, carretera a San Francisco, 1000–1500 m, 13 abr 1991, Giraldo-Cañas 71 (HUA).

Serpocaulon catharinae (Langsd. et Fisch.) A. R. Sm., Brasil, Paraná: Curitiba, campus Politécnico Universidad Federal de Paraná, frente a la Facultad de Ciencias Biológicas, 23 June 2010, Sanín 4199 (HUA).

Serpocaulon crystalloneuron (Rosenst.) A. R. Sm., Bolivia, Provincia de Nor Yungas: Municipio de Coroico, Parque Nacional Cota Pata, Camino antiguo de Sillutinkara. 14 January 2012, Sanín et al. 5530 (HUA).

Serpocaulon dasypleuron (Kunze) A. R. Sm., Colombia, Amazonas: río Miriparaná, Cerro de la Gente Chiquita in head waters of Quebrada Guacayá, sin dat., 18 May 1952, Schultes 16501 (COL).

Serpocaulon dissimile (L.) A. R. Sm., Costa Rica, San Vito de Coto Brus: Jardín Botánico Robert y Catherine Wilson, Estación Biológica Las Cruces, O.E.T., sendero vía al Río Java, 1211 m, N 08º47ʹ0.3‘, W 82º57ʹ34’, 14 June 2011, Sanín & Santiago 5019 (HUA).

Serpocaulon eleutherophlebium (Fée) A. R. Sm., Colombia, Cauca, Popayán, Parque Nacional Natural Puracé, sector anexo a las bases militares, 3887 m, 02° 22ʹ N, 76° 26ʹ W, 26 July 2009, Sanín & Sanín 3293 (FAUC).

Serpocaulon falcaria (Kunze) A. R. Sm., Costa Rica; Cordillera de Talamanca; Cerro de la Muerte, al margen de la carretera vía Savegre, 2484 m, 09° 32ʹ N, 83° 48ʹ W, 20 June 2011, Sanín & Torrez 5031 (HUA).

Serpocaulon fraxinifolium (Jacq.) A. R. Sm., Bolivia, Provincia de Nor Yungas: Municipio de Coroico, Parque Nacional Cota Pata, Camino antiguo de Sillutinkara, 2444 m, 16° 15ʹ N, 67° 53ʹ W, 15 January 2012, Sanín et al. 5537 (HUA).

Serpocaulon intricatum (M. Kessler et A. R. Sm.) A. R. Sm., Bolivia, Provincia de Nor Yungas: Municipio de Coroico, Parque Nacional Cota Pata, Camino antiguo de Sillutinkara, 2178 m, 16° 14ʹ N, 67° 53ʹ W, 15 January 2012, Sanín et al. 5526 (HUA).

Serpocaulon lasiopus (Klotzsch) A. R. Sm., Colombia, Cundinamarca: Laguna de Ubaque, 2000 m, 11 July 1974, Acosta-Arteaga 595 (COL).

Serpocaulon latipes (Langsd. et L. Fisch.) A. R. Sm., Brazil, Paraná: Curitiba, Paranaguá, río Guaraguaçu, Entrada para Sambaqui, 11 m, 25° 39ʹ S, 48° 30ʹ W, 1 July 2010, Sanín & Labiak 4200 (HUA).

Serpocaulon levigatum (Cav.) A. R. Sm., Colombia, Caldas: Manizales-Villamaria, vereda Montaño, finca San Antonio, relicto al nor-oriente de la casa, 2226 m, 4° 59ʹ N, 75° 27ʹ W, 7 Ago 2009, Sanín et al. 3315 (FAUC).

Serpocaulon loriceum (L.) A. R. Sm., Costa Rica, San Vito de Coto Brus: Jardín Botánico Robert y Catherine Wilson, Estación Biológica Las Cruces, O.E.T., sendero vía al Río Java, Sanín & Santiago 5014 (HUA).

Serpocaulon maritimum (Hieron.) A. R. Sm., Colombia, Nariño: Junín, en dirección a Barbacoas, 01° 20ʹ N, 78° 02ʹ W, 1000 m, 20 Dic 1972, Haegemann & Leist 1750 (COL).

Serpocaulon menisciifolium (Langsd. et Fisch.) A. R. Sm., Brasil, Paraná: Antonina, Reserva Natural Rio Cachoeira (SPVS), estrada Sede-Gervásio, Matos 544 (UPCB).

Serpocaulon patentissimum (Mett. ex Kuhn) A. R. Sm., Colombia, Nariño: Reserva Natural la Planada, sendero el Rondón cerca 2 km del Centro de Investigación, 1900 m, 01° 09ʹ N, 77° 58ʹ W, 19 January 1997, Herrera 9140 (PSO).

Serpocaulon polystichum (Link) A. R. Sm., Colombia, Antioquia: Valdivia, vereda San Fermín, 1 km de la vía Ventanas (Mun. Yarumal) Briceño, 1700 m, 07° 08ʹ N, 75° 30ʹ W, 22 March 1988, Arbeláez et al. 313 (HUA).

Serpocaulon ptilorhizon (Christ) A. R. Sm., Colombia, Antioquia: Guatapé, vereda Santa Rita, finca Montepinar, bosque pluvial, 1850 m, 06° 16ʹ N, 75° 10ʹ W, 6 February 1990, Echeverry 195 (HUA).

Serpocaulon richardii (Klotzsch) A. R. Sm., Colombia, Boyacá: carretera a Pauna-Otro Mundo, 1350 m, 16 October 1967, Jaramillo-Mejía et al. 3650 (COL).

Serpocaulon sessilifolium (Desv.) A. R. Sm., Bolivia, Provincia de Nor Yungas: Coroico, Parque Nacional Cota Pata, Camino antiguo de Sillutinkara, 1962 m, 16° 13ʹ S, 67° 52ʹ W, 15 January 2012, Sanín et al. 5534 (HUA).

Serpocaulon subandinum (Sodiro) A.R. Sm., Bolivia, Provincia de Nor Yungas Municipio de Coroico, Parque Nacional Cota Pata, Camino Precolombino de Sillutinkara, 3069 m, 16° 16ʹ S, 67° 53ʹ W, 14 January 2012, Sanín et al. 5521 (HUA).

Serpocaulon triseriale (Sw.) A. R. Sm., Colombia, Caldas: Neira, vía Aránzazu, 4 km del casco urbano, 1350 m, 05° 12ʹ N, 75° 35ʹ W, 3 March 2008, Sanín 2602 (FAUC).

Serpocaulon wagneri (Mett.) A. R. Sm., Colombia, Cundinamarca: Ubalá, Vereda San Roque, camino a Campo Hermoso, 1150 m, 30 June 1998, Fernández-Alonso 16217 (COL).

Campyloneurum brevifolium (Lodd. ex Link) Link. Colombia, Antioquia, Anorí, Región Providencia, sector Aljibes-La Tirana, 500 m, 18 January 2004, Rodríguez 4484 (COL).

Microgramma lycopodioides (L.) Copel. Colombia, Norte de Santander, San Cayetano, Vereda Ayacucho, Finca Bellavista, 1355 m, 11 May 2009, Infante-Betancour 623 (COL).

Pleopeltis bombycina (Maxon) A.R. Sm. Colombia, Antioquia, Municipio de San Rafael, Morro Pan de Azúcar, 1100–1354 m, 14 October 1981, Orozco, et al. 754 (COL).

Polypodium vulgare L. Alemania, Poiana, Uzului, 500 m, 22 August 1970, Barabas & Mititelu 1085 (COL).