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Spore morphology of the Mexican species of the Elaphoglossum petiolatum complex (Dryopteridaceae)

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Abstract

Elaphoglossum is a taxonomically challenging fern genus with about 600 species. The present study concerns a group of five Elaphoglossum species mainly distributed in Mexico, four of which were recovered as monophyletic and are here called the Mexican-petiolatum clade (E. petiolatum, E. potosianum, E. pringlei, and E. rzedowskii). The fifth species, E. muelleri, is distributed in Central America and Mexico and is morphologically similar to species of the Mexican-petiolatum clade. This study investigates the spores of these five species and has three objectives: first, to describe and compare their spore ornamentation; second, to analyze the variation in spore size among the species and within the widely distributed species E. petiolatum; and third, to assess whether perine ornamentation and exine size are useful to recognize species of the Mexican-petiolatum clade and/or for species delimitation. We studied 26 herbarium specimens of the five species and used scanning electron microscopy and statistical analyses to examine, measure, and analyze 133 spores. All the spores examined have uniform primary ornamentation (broad and discontinuous folds), but the secondary ornamentation is variable. Spores from Mexican specimens, regardless of their species, have small spines with narrow bases; spores of E. petiolatum from Central America have spines with wider bases; and spores of E. petiolatum from the West Indies have verrucae. Statistical analyses revealed that there are significant differences in spore size among all the studied species and within specimens identified as E. petiolatum. Within E. petiolatum, spores from the Dominican Republic are larger than those from the other countries. The results support our hypothesis that E. muelleri is part of the Mexican-petiolatum clade and suggest that E. petiolatum in Mexico, Central America, and the West Indies likely represent different taxa. Spore size differences help distinguish closely related species in the group such as E. pringlei and E. rzedowskii and indicate that there might be polyploid species within the Mexican-petiolatum clade.

Resumen

Elaphoglossum es un género de helechos taxonómicamente desafiante con alrededor de 600 especies. El presente estudio se enfocó en un grupo de cinco especies pertenecientes al género Elaphoglossum, distribuidas principalmente en México. De éstas, cuatro se recuperaron en un grupo monofilético denominado aquí como el clado mexicano de E. petiolatum (E. petiolatum, E. potosianum, E. pringlei y E. rzedowskii). La quinta especie, E. muelleri, se distribuye en América Central y México y es morfológicamente similar a las especies del clado mexicano de E. petiolatum. Este estudio investiga las esporas de estas cinco especies y tiene tres objetivos: primero, describir y comparar la ornamentación de las esporas; segundo, analizar la variación en el tamaño de las esporas entre las especies y dentro de la especie ampliamente distribuida E. petiolatum; y tercero, evaluar si la ornamentación de la perina y el tamaño de la exina son útiles para reconocer y/o delimitar estas especies. Examinamos, medimos y analizamos 133 esporas de 26 especímenes de herbario pertenecientes a las cinco especies, mediante microscopía electrónica de barrido y análisis estadísticos. Todas las esporas examinadas tienen ornamentación primaria uniforme (pliegues anchos y discontinuos), pero la ornamentación secundaria es variable. Las esporas de especímenes mexicanos, independientemente de su especie, tienen espinas pequeñas con bases estrechas; las esporas de E. petiolatum de América Central tienen espinas con bases más anchas; y las esporas de E. petiolatum de las Antillas tienen verrugas. Los análisis estadísticos revelaron que existen diferencias significativas en el tamaño de las esporas entre todas las especies estudiadas y dentro de los especímenes identificados como E. petiolatum. Dentro de E. petiolatum, las esporas de República Dominicana son más grandes que las de los demás países. Los resultados obtenidos apoyan nuestra hipótesis de que E. muelleri es parte del clado mexicano de E. petiolatum y sugieren que E. petiolatum de México, América Central y las Antillas, probablemente representan taxones diferentes. Las diferencias en el tamaño de las esporas ayudan a distinguir especies estrechamente relacionadas como E. pringlei y E. rzedowskii, e indican que puede haber especies poliploides dentro del clado mexicano de E. petiolatum.

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Acknowledgments

This work is part of the Ph. D thesis of the first author at the Posgrado en Ciencias Biológicas (PCB) of the Universidad Nacional Autónoma de México (UNAM), supported by the Consejo Nacional de Ciencia y Tecnología (CONACyT) with a Ph.D. fellowship (508729), under the advice of A. Vasco and E. Solano. This study was supported in part by “Programa de Apoyo a Proyectos de Investigación e Innovación Tecnológica” (PAPIIT), Universidad Nacional Autónoma de México (IA201416 to A.Vasco). Fieldwork was partially funded by a “Research Grant for Graduate Students” from the American Society of Plant Taxonomists (ASPT, 2018 to A.G. Martínez-Bercerril). We are grateful to the curators and collection managers of the herbaria FEZA, MEXU (especially Laura Calvillo-Canadell, Angélica Ramírez Roa, and María del Rosario García Peña), NY, and MO for specimen loans and for allowing us to sample spores for SEM imaging. We thank all the people who accompanied us to collect specimens and to the communities that allowed us to collect in their lands. We are grateful to Berenit Mendoza Garfias from the Microscopy and Biodiversity Photograph Laboratory (Instituto de Biología UNAM) for help with taking spore SEM images. We appreciate the discussions and advise of Drs. Teresa M. Terrazas, David S. Gernandt, and Daniel Tejero Diez. Special thanks to Robbin Moran (New York Botanical Garden) for his reviews and for letting us use his SEM images posted on plantsystematics.org. We appreciate the comments and careful reviews from two anonymous reviewers.

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Authors and Affiliations

  1. Facultad de Estudios Superiores Zaragoza, Unidad de Investigación en Sistemática Vegetal y Suelo, Universidad Nacional Autónoma de México, Batalla 5 de mayo s/n, Ejército de Oriente, Iztapalapa, C.P. 09230, Ciudad de México, México

    Ana Gabriela Martínez-Becerril & Eloy Solano

  2. Posgrado en Ciencias Biológicas, Universidad Nacional Autónoma de México, Ciudad de México, México

    Ana Gabriela Martínez-Becerril

  3. Botanical Research Institute of Texas, 1700 University Drive, Fort Worth, TX, 76107-3400, USA

    Alejandra Vasco

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  1. Ana Gabriela Martínez-Becerril
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  2. Eloy Solano
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  3. Alejandra Vasco
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Correspondence to Alejandra Vasco.

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Appendix 1. Voucher information for all Elaphoglossum spores included in the analyses. Stars indicate spore images taken from Plantsystematics.org

Appendix 1. Voucher information for all Elaphoglossum spores included in the analyses. Stars indicate spore images taken from Plantsystematics.org

Elaphoglossum muelleri (E. Fourn.) C. Chr. MEXICO. Anderson 13,211 (MEXU); McVaugh 23,885 (MEXU). Elaphoglossum petiolatum (Sw.) Urb. COSTA RICA. Lloyd 4134 (NY); Moran 3079* (NY). DOMINICAN REPUBLIC. Garcia 958 (NY); Lloyd 753* (NY). MEXICO. Gallardo-Hernandez 836 (MEXU); Reveal & Atwood 3572* (NY). JAMAICA. Proctor 5297 (MO); Underwood 545, 3141*, 3214 (NY). PANAMA. Croat 26,996 (NY). Elaphoglossum potosianum Christ. MEXICO. Carranza 5140 (MEXU, NY*), 2036 (MEXU); Martínez-Becerril 399 (FEZA, MEXU); Parry & Palmer 1007* (NY). Elaphoglossum pringlei (Davenp.) C. Chr. MEXICO. Anderson 4778 (MEXU); Ernest 2371 (MEXU); Martínez-Becerril 405, 407 (FEZA, MEXU); Mickel 4463* (NY). Elaphoglossum rzedowskii Mickel. MEXICO. Cornejo-Tenorio 1438 (MEXU); Martínez-Becerril 406, 408 (FEZA, MEXU); Rzedowski 26032* (NY).

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Martínez-Becerril, A.G., Solano, E. & Vasco, A. Spore morphology of the Mexican species of the Elaphoglossum petiolatum complex (Dryopteridaceae). Brittonia 73, 131–142 (2021). https://doi.org/10.1007/s12228-020-09643-8

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