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Phylogenetics and historical biogeography of Encyclia (Laeliinae: Orchidaceae) with an emphasis on the E. adenocarpos complex, a new species, and a preliminary species list for the genus

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Abstract

We explore the phylogenetic position of the Encyclia adenocarpos complex through a multilocus  analysis of Encyclia with the following DNA regions: ITS and plastid rpl32-trnL, trnL-F, and ycf1, analyzed under the Bayesian inference and Maximum Parsimony paradigms. We also performed an analysis of reconstruction of ancestral areas, with particular interest in the first diverging lineages of Encyclia. We used an ad hoc regionalization system designed to fit our distributional Encyclia data to reconstruct the ancestral distribution of the genus and the most relevant nodes. The analyses yielded a moderately well-supported topology with the broadest taxonomic Encyclia sampling yet. Our results indicate the Encyclia adenocarpos complex is monophyletic, highly supported, and sister to a large clade with ca. 95% of the species of Encyclia included in the analyses and suggest the clade is composed of several related, fundamentally allopatric species distributed along the Pacific slopes of Megamexico, including a novelty here proposed, Encyclia mariaeugeniae, which is closely related to Encyclia enriquearcilae yet differing in several diagnostic characters, such as a narrower central lobe to the labellum. The novelty was assessed as EN under IUCN criteria. We provide a key to the E. adenocarpos clade. Our Encyclia phylogeny identifies several clades displaying strong geographic signal; some of these are discussed in terms of morphology, ecological preferences, and pollination syndromes. The reconstruction of ancestral areas indicates with high probability that the earliest diverging nodes of Encyclia occurred in Megamexico. Floral variation within clades suggests the genus colonized geographical areas and underwent diversifications to occupy novel pollination syndromes in a pattern of allopatric, similar assemblages of syndromes composed of unrelated taxa. We present a preliminary species list of Encyclia and their distributions along the major biogeographic areas to provide a hypothesis of diversity patterns and the biogeographical areas where the species occur. As we currently understand the genus, Encyclia consists of 213 taxa, including 179 formally proposed species, 10 undescribed ones, and 25 nothotaxa. We provide five plates depicting 100 species to document the morphological and geographical diversity of Encyclia.

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Data availability

The datasets generated and/or analyzed during the current study are available from the corresponding author on reasonable request.

Change history

  • 10 September 2022

    Supplementary Table S1 has been added.

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Acknowledgements

The authors would like to acknowledge Rodrigo Duno de Stefano for comments on a previous version of this manuscript and subsequent discussion. Two anonymous reviewers and the editor of the journal contributed commentary and suggestions for which we are deeply indebted and that made this a much better article. Carlos García Esquivel, Ramón López, David Hunt, Eduardo A. Pérez García, Patricia Harding, Dennis Szeszko, and Alejandro Zabalgoitia commented earlier drafts of the manuscript and/or provided crucial input on circumscriptions and distributional issues of the genus, as well as photographs. Bosco Javier Zambrano Romero, of Orquiecuador & Gloxinias contributed commentary on Ecuadorean Encyclia and allowed us to use some of his good photographs. Manuel Calvo provided discussion and materials on Cuban Encyclia for which we are deeply indebted. Gerardo Castiglione (Mérida, Venezuela), Saul Hernández (Morelia, Mexico) contributed photographs and comments on Encyclia. Anderson Cassano, from Sao Paulo, Brazil contributed many good photographs of Brazilian species, for which we are deeply indebted. Manfred Speckmaier of the Vienna Botanical Gardens contributed photographs. Luis Ocupa Horna contributed photographs of Peruvian Encyclia species. Carlos L. Leopardi Verde (University of Colima) contributed photographs and discussion. Julio Valdez Partida (Universidad Michoacana de San Nicolás de Hidalgo) shared his knowledge and photographs of the Encyclia meliosma/E. spatella complex for which we are deeply indebted. Enrique Arcila Arcila, from Mérida, Yucatán, Mexico, cultivated and allowed access to many Encyclia plants that were crucial for our understanding of the species of the genus. Peter W. Fritsch at BRIT allowed us the use of a photograph of E. yauaperyensis. Fredy Archila allowed us to use a photograph of E. delacruzii for which we are deeply grateful. Francis Yuliett Echeverría Bobadilla allowed us to use some of her photographs. Teague Embrey was of great help in a field trip of one of us (ITC) in search of Encyclia mariaeugeniae. María Eugenia Cen Alonzo helped cultivating species of Encyclia. Verónica Ortega Baranda, Deysi Lizeth de la Cruz Salinas, and María de los Ángeles Luis Reyna of Jardín Botánico de Puerto Escondido provided crucial information on E. mariaeugeniae. We thank the curators of AMO, BIGU, CHIP, COL, ENC, F, FCME, HEM, HUT, IBUG, K, M, MEXU, MO, NY, OAX, RENZ, SEL, SERO, TEFH, UAMIZ, US, VEN, and W for allowing us access their collections. Silvia Hernández-Aguilar (CICY) assisted with the management of herbarium specimens. Néstor E. Raigoza (CICY) helped in the laboratory. GC thanks The Marie Selby Botanical Gardens for a research scholarship in May 2017 and June 2019 that were instrumental to the research, results of which are presented herein. Günter Gerlach hosted GC during a research visit at M in 2012. Ernst Vitek, Manfred Speckmaier, and Bruno Wällnofer were of great help for GC during his 2012 visit to W (Naturhistorisches Museum Wien, Austria). DAAD (the German Academic Exchange Service) provided a research scholarship for GC in 2012 when type and other relevant Encyclia materials were studied. Kanchi Gandhi (HUH) assisted us in nomenclatural matters. IMR would like to acknowledge the logistical support of Pablo Carrillo Reyes, Claudia Ramírez, and Alejandro Zabalgoitia while conducting field work in Jalisco and neighboring states. A photograph of E. adenocarpos was borrowed from the SEINet Portal Network. 2020. http//:swbiodiversity.org/seinet/index.php., Accessed on September 21 while another photograph of Alex Popovkin of E. oncidioides comes from https://commons.wikimedia.org/wiki/File:Encyclia_oncidioides_(Lindl.)_Schltr._-_Flickr_-_Alex_Popovkin,_Bahia,_Brazil_(7).jpg.

Funding

CONACyT partially funded this project via grant 68640 “Sistemática y filogenia del género Encyclia Hook. con énfasis en Megaméxico”, awarded to GC and a graduate scholarship to ITC (CVU 568712). The American Orchid Society also partially funded this project via the research grant adjudicated to the project “Systematics and evolution of Encyclia Hook. s.s. (Orchidaceae: Laeliinae) with emphasis in Megamexico” awarded to GC. GAR-G acknowledges the financial support of the Orchid Society of Arizona.

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Authors and Affiliations

  1. Herbarium CICY, Centro de Investigación Científica de Yucatán, Colonia Chuburná de Hidalgo, Mérida, A.C., Calle 43 #130 x 32 y 34, Yucatán, C.P. 97205, México

    Germán Carnevali, Iván Tamayo-Cen, Ivón M. Ramírez‑Morillo, José L. Tapia-Muñoz & Gustavo A. Romero-González

  2. Orchid Herbarium of Oakes Ames, Harvard University Herbaria, 22 Divinity Avenue, MB, 02138, Cambridge, U.S.A.

    Germán Carnevali & Gustavo A. Romero-González

  3. UMAR, Universidad del Mar Campus Puerto Escondido, San Pedro Mixtepec, Juquila, Oaxaca, C.P. 71980, México

    Carlos E. Méndez-Luna

  4. Tecnológico Nacional de México, Instituto Tecnológico de Chiná, Calle 11, Col. Centro Chiná, Campeche, C.P. 24050, México

    William Cetzal-Ix

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Carnevali, G., Tamayo-Cen, I., Méndez-Luna, C.E. et al. Phylogenetics and historical biogeography of Encyclia (Laeliinae: Orchidaceae) with an emphasis on the E. adenocarpos complex, a new species, and a preliminary species list for the genus. Org Divers Evol 23, 41–75 (2023). https://doi.org/10.1007/s13127-022-00575-7

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