A nomenclator for Elaphoglossum section Polytrichia (Dryopteridaceae), with notes on the identification and biogeography of its species

Fernando Matos

Elaphoglossum section Polytrichia is one of the main clades of the fern genus Elaphoglossum. It is usually distinguished from the rest of the genus by the presence of subulate scales and lack of hydathodes. Here we provide a nomenclatural synopsis with notes on the species taxonomy and identification, along with comments on the clade’s biogeography. A total of 80 names were found to apply to species in the Polytrichia clade. Based on an examination of 3475 herbarium gatherings, we recognize in the group 53 species and two hybrids. The clade is almost entirely Neotropical, occurring from Mexico and the Antilles to Argentina and Brazil. The only exception is E. hybridum, which is widespread in South America but also occurs in Africa, Madagascar, and islands of the mid-Atlantic and Indian Oceans. We provide maps of geographic distribution and species richness for the section. Each species is provided with place and date of publication, information on types, complete synonymy, notes on geographic distribution, and other pertinent remarks. The following nomenclatural combinations are made: E. versatile and E. occidentale, comb. nov. Also, one epitype is designated for E. clathratum, lectotypes are designated for 13 names (Acrostichum barbatum, A. hybridum var. minor, A. mexicanum, A. nitidum, A. stramineum, E. angustioblongum, E. cinctum, E. cordifolium, E. demissum, E. erinaceum var. boliviense, E. lindbergii var. truncatum, E. reductum, E. truncatum), and neotypes are designated for E. hystrix and E. versatile. A list of the specimens examined is provided.

Phytotaxa 535 (1): 001–067 https://www.mapress.com/pt/ Copyright © 2022 Magnolia Press ISSN 1179-3155 (print edition) Monograph PHYTOTAXA ISSN 1179-3163 (online edition) https://doi.org/10.11646/phytotaxa.535.1.1 PHYTOTAXA 535 A nomenclator for Elaphoglossum section Polytrichia (Dryopteridaceae), with notes on the identification and biogeography of its species FERNANDO B. MATOS1,2,4, ROBBIN C. MORAN2,5 & JEFFERSON PRADO3,6 1 Universidade Federal de Viçosa, Departamento de Biologia Vegetal, Av. Purdue s.n., CEP 36570-900, Viçosa, MG, Brazil 2 The New York Botanical Garden, Institute of Systematic Botany, 2900 Southern Blvd., Bronx, NY 10458, U.S.A. 3 Instituto de Pesquisas Ambientais (IPA), Av. Miguel Estéfano, 3687 CEP 04301-902, São Paulo, SP, Brasil 4 � fbtms@yahoo.com.br; https://orcid.org/0000-0003-2133-6032 5 � rmoran@nybg.org; https://orcid.org/0000-0002-9356-8265 6 � jprado.01@uol.com.br; https://orcid.org/0000-0003-4783-3125 Corresponding author: Fernando B. Matos, � fbtms@yahoo.com.br Magnolia Press Auckland, New Zealand Accepted by Marcus Lehnert: 4 Dec. 2021; published: 21 Feb. 2022 FERNANDO B. MATOS, ROBBIN C. MORAN & JEFFERSON PRADO A nomenclator for Elaphoglossum section Polytrichia (Dryopteridaceae), with notes on the identification and biogeography of its species (Phytotaxa 535) 67 pp.; 30 cm. 21 February 2022 ISBN 978-1-77688-460-5 (paperback) ISBN 978-1-77688-461-2 (Online edition) FIRST PUBLISHED IN 2022 BY Magnolia Press P.O. Box 41-383 Auckland 1041 New Zealand e-mail: magnolia@mapress.com https://www.mapress.com/pt/ © 2022 Magnolia Press All rights reserved. No part of this publication may be reproduced, stored, transmitted or disseminated, in any form, or by any means, without prior written permission from the publisher, to whom all requests to reproduce copyright material should be directed in writing. This authorization does not extend to any other kind of copying, by any means, in any form, and for any purpose other than private research use. ISSN 1179-3155 (print edition) ISSN 1179-3163 (online edition) • Phytotaxa 535 (1) © 2022 Magnolia Press MATOS ET AL. Abstract Elaphoglossum section Polytrichia is one of the main clades of the fern genus Elaphoglossum. It is usually distinguished from the rest of the genus by the presence of subulate scales and lack of hydathodes. Here we provide a nomenclatural synopsis with notes on the species taxonomy and identification, along with comments on the clade’s biogeography. A total of 80 names were found to apply to species in the Polytrichia clade. Based on an examination of 3475 herbarium gatherings, we recognize in the group 53 species and two hybrids. The clade is almost entirely Neotropical, occurring from Mexico and the Antilles to Argentina and Brazil. The only exception is E. hybridum, which is widespread in South America but also occurs in Africa, Madagascar, and islands of the mid-Atlantic and Indian Oceans. We provide maps of geographic distribution and species richness for the section. Each species is provided with place and date of publication, information on types, complete synonymy, notes on geographic distribution, and other pertinent remarks. The following nomenclatural combinations are made: E. versatile and E. occidentale, comb. nov. Also, one epitype is designated for E. clathratum, lectotypes are designated for 13 names (Acrostichum barbatum, A. hybridum var. minor, A. mexicanum, A. nitidum, A. stramineum, E. angustioblongum, E. cinctum, E. cordifolium, E. demissum, E. erinaceum var. boliviense, E. lindbergii var. truncatum, E. reductum, E. truncatum), and neotypes are designated for E. hystrix and E. versatile. A list of the specimens examined is provided. Keywords: ferns, hydathodes, nomenclature, subulates, taxonomy, typification Introduction Elaphoglossum Schott ex J.Sm. is the largest genus in the Dryopteridaceae, which is the largest family of ferns (PPG I 2016). Estimates of the number of species worldwide in the genus range from 600 to 800 (Mickel & Atehortúa 1980, Mabberley 2008, Rouhan 2020). Within the Dryopteridaceae, Elaphoglossum belongs to the subfamily Elaphoglossoideae (Pic.Serm.) Crabbe, and within the subfamily to the bolbitidoid clade, which also contains five other genera: Arthrobotrya, Bolbitis, Lomagramma, Mickelia, and Teratophyllum. The bolbitidoids are characterized by the synapomorphies of dorsiventral stems with a transversely elongated ventral meristele, ventral root insertion, strongly dimorphic sterile-fertile leaves, and acrostichoid sori (Moran et al. 2010a). Within the bolbitidoids, Elaphoglossum is monophyletic and sister to Mickelia R.C.Moran, Labiak & Sundue (Rouhan et al. 2004, Schuettpelz & Pryer 2007, Moran et al. 2010a, 2010b, Liu et al. 2016, PPG I 2016). Elaphoglossum is distinguished from Mickelia by simple and entire leaves, free veins, and conspicuous phyllopodia (Holttum 1978, Mickel & Atehortúa 1980, Moran et al. 2010a). There are rare exceptions to some of these characters, such as a few species with anastomosing veins (e.g., E. crinitum (L.) Christ, E. crassifolium (Gaudich.) W.R.Anderson & Crosby, and E. pachyphyllum (Kunze) C.Chr.), and six species with leaves either pedate, pinnate, or flabellate (Wagner 1954, Vasco et al. 2009, 2013). Several authors have attempted to divide Elaphoglossum into smaller groups on the basis of morphology (e.g., Fée 1845, 1852, Moore 1857–1862, Sodiro 1893, Christ 1899, Diels 1902, Brade 1961, Holttum 1978, Mickel & Atehortúa 1980). More recently, the phylogenetic relationships within Elaphoglossum have been examined using plastid DNA sequences (e.g., Rouhan et al. 2004, Skog et al. 2004, Lóriga et al. 2014, Vasco et al. 2015, Matos et al. 2019). One of the main clades that has always been recovered by these studies is the “subulate-scaled clade” (sensu Skog et al. 2004), which is characterized by the presence of patent, hairlike scales with revolute margins. It is now generally agreed that this clade comprises two major subclades distinguished by the presence versus absence of hydathodes (e.g., Rouhan et al. 2004, Lóriga et al. 2014, Matos et al. 2019). The hydathodous clade corresponds to sects. Eximia Mickel & Atehortúa, Setosa (Christ) Mickel & Atehortúa, and Undulata Christ, as defined by Mickel & Atehortúa (1980). The non-hydathodous clade corresponds to sections Decorata Mickel & Atehortúa and Polytrichia Mickel & Atehortúa (1980). These two non-hydathodous sections, which have never received comprehensive monographic study, are the focus of the present paper (Fig. 1). Taxonomic history of Elaphoglossum sect. Polytrichia (Sodiro) Christ The oldest name in Elaphoglossum sect. Polytrichia is Acrostichum crinitum L. (Linnaeus 1753), which was described in a time when Acrostichum L. included all ferns with acrostichoid sori (Mickel & Atehortúa 1980). Its original description was based on an illustration by Petiver (1712, tab. 13, f. 14), which was copied from Plumier (1705, tab. 125) and represented an unvouchered plant from Martinique (see Matos et al. 2015). This species, which is now A NOMENCLATOR FOR ELAPhoGLoSSUM SECTION PoLYTrIChIA Phytotaxa 535 (1) © 2022 Magnolia Press • 3 recognized as E. crinitum, is easily distinguished by its large leaves with broadly elliptic laminae, reticulate veins, and dark subulate scales (Figure 2D, 3I, 4A–B). Within Elaphoglossum, reticulate venation is rare, partially explaining why this species has been placed in different genera, such as olfersia Raddi (Presl 1836), hymenodium Fée (Fée 1845), Dictyoglossum Sm. (Smith 1846), Anetium Splitg. (Presl 1851), and Chrysodium Fée (Mettenius 1856). The species was finally transferred to Elaphoglossum by Christ (1899). Since then, Christ’s opinion has been supported by morphological (e.g., Copeland 1947, Mickel & Atehortúa 1980) and molecular phylogenetic studies (e.g., Rouhan et al. 2004, Skog et al. 2004, Schuettpelz & Pryer 2007, Matos et al. 2019). FIGURE . Consensus phylogeny depicting the relationships among major clades of Elaphoglossum. Topology summarizes results of previously published phylogenetic studies (Skog et al. 2004; Rouhan et al. 2004; Lóriga et al. 2014; Vasco et al. 2015; Matos et al. 2019). The Polytrichia Clade, which is the focus of the present study, is indicated in black. The asterisk indicates the presence of hydathodes in E. amygdalifolium. Fée (1845) was the first to provide a monograph for the acrostichoid ferns. He created hymenodium to accommodate Elaphoglossum crinitum (along with h. crassifolium Fée and h. kunzeanum Fée) and included eight other names that are now recognized in E. sect. Polytrichia under Acrostichum: A. apodum Kaulf., A. decoratum Kunze, A. erinaceum Fée, A. hybridum Bory, A. hystrix Kunze, A. platyneuron Fée, and A. scolopendrifolium Raddi. Sodiro (1893) was the first to use the name Polytrichia, in a tentative division of the genus for Ecuador. In his dichotomous key, he grouped all species with subulate scales under the infrageneric name Setosa. Based on indument • Phytotaxa 535 (1) © 2022 Magnolia Press MATOS ET AL. abundance, he further divided the subulate-scaled species into two unranked groups: oligotrichia (scales few) and Polytrichia (scales many). According to the Code (Turland et al. 2018), these two names are validly published, even though Sodiro (1893) did not provide a clear indication of their ranks (see Turland et al. 2018, Art. 37.3) or indicate types (see Turland et al. 2018, Art. 40.1). It is also of interest that Sodiro (1893) placed all subulate-scaled species without hydathodes under Acrostichum [infragen. unranked] oligotrichia, with the exception of A. tambillense, which was placed in a group comprised of glabrous species (i.e., Acrostichum [infragen. unranked] Glabra). FIGURE . Field photos of selected species of Elaphoglossum sect. Polytrichia. A. E. angustioblongum. B. E. backhouseanum, showing nest-forming leaves. C. E. cotoi. D. E. crinitum. E. E. decoratum. F. Two of us authors (F. Matos and R. Moran) with E. erinaceum in Costa Rica. G. E. glaziovii. H. E. miersii, growing on the trunk of a tree fern. I. E. procurrens. A–C by F. Matos in Costa Rica; D by M. Sundue in Costa Rica; E by R. Moran in Costa Rica; F by M. Sundue; G–H by F. Matos in Brazil; I by J. Lóriga in Cuba. A NOMENCLATOR FOR ELAPhoGLoSSUM SECTION PoLYTrIChIA Phytotaxa 535 (1) © 2022 Magnolia Press • 5 A few years later, in his monograph of Elaphoglossum, Christ (1899) provided a short description in German and assigned sectional rank to Polytrichia. The name oligotrichia was not considered by Christ (1899), and the subulatescaled species without hydathodes were placed into four different groups: “divisio Melanolepidea”, “divisio Decorata”, and three subgroups of “sectio Polytrichia” (i.e., hybrida, hymenodium, and Pilosella). Mickel & Atehortúa (1980) proposed a new infrageneric classification for Elaphoglossum based on morphological features. They created section Decorata Mickel & Atehortúa to accommodate the “remarkably distinct and without close relatives” E. decoratum (Kunze) T.Moore. They also placed all subulate-scaled species without hydathodes in section Polytrichia (Sodiro) Christ, designated Acrostichum crinitum as the lectotype of this section, and further divided its species into three groups. The first of these groups was subsect. hymenodium (Fée) Christ, with a single species (Elaphoglossum crinitum) characterized by broadly elliptic leaves and reticulate veins. The second group was subsect. Apoda Mickel & Atehortúa, with three species (E. apodum (Kaulf.) Schott ex J.Sm., E. cubense (Mett. ex Kuhn) C.Chr., and E. siliquoides (Jenman) C.Chr.) characterized by nearly sessile leaves and light-colored (orange to brown) subulate scales evenly distributed on the laminae. Finally, the third and largest group was subsect. hybrida Christ, with an unknown number of species characterized by long petioles and subulate scales usually limited to petioles, costae, and laminar margins. Mickel & Atehortúa (1980) listed 12 names under this subsection. A few years later, Atehortúa (1983) studied subsect. Apoda for her doctoral thesis, which resulted in two papers about the anatomy, morphology, and phytochemistry of these plants (Atehortúa 1984, 1985). In her thesis, she removed Elaphoglossum siliquoides to E. sect. Setosa subsect. Pilosella Christ and recognized nine taxa in a group informally called the “Elaphoglossum apodum complex”. The taxa she included in this group were E. amazonicum Atehortúa, E. apodum, E. auripilum Christ var. auripilum, E. auripilum var. longipilosum Atehortúa, E. backouseanum T.Moore, E. cubense, E. latum (Mickel) Atehortúa, E. procurrens (Mett. ex D.C.Eaton) T.Moore, and E. raywaense (Jenman) Alston. The three new names introduced by Atehortúa (1983), however, were not validly published because her doctoral thesis does not constitute effective publication (Turland et al. 2018, Art. 30.9). These names were later validated by Mickel (1991), as E. amazonicum Atehortúa ex Mickel, and Mickel (1992), as E. auripilum var. longipilosum Atehortúa ex Mickel and E. latum (Mickel) Atehortúa ex Mickel. Although Elaphoglossum sect. Polytrichia has never been fully monographed, many of its species have been included in checklists and floras from different regions of the world (e.g., Alston 1958, Proctor 1977, 1985, 1989, Mickel 1981, 1985, 1991, 1995a, 1995b, Lorence & Rouhan 2004, Mickel & Smith 2004). Most of these studies have not treated the species of sect. Polytrichia separately, albeit these species usually key out together because of the presence of subulate scales and absence of hydathodes. Three recent floras have divided the genus in sections and provided a separate key for sect. Polytrichia: Kessler et al. (2018, for Bolivia), Matos (2020a, for Brazil), and Rouhan (2020, for Madagascar). Two other studies have focused exclusively on E. sect. Polytrichia: Rojas-Alvarado (2003, for Mexico and Central America) and Matos & Mickel (2014, for Brazil). In total, about 30 new taxa have been published since Mickel & Atehortúa (1980) recircumscribed the section (e.g., Lorence 1984, Mickel 1987, 1991, 1992, Rojas-Alvarado 1996a, 2002, 2003, 2009, 2010, 2017, Mickel & Smith 2004, Moran & Mickel 2004, Matos & Moran 2013, 2017, Matos & Mickel 2014, Matos et al. 2019, Matos et al. 2021), including one variety (E. erinaceum var. occidentale Mickel) and two nothospecies: E. × setaceum Lorence (Lorence 1984) and E. × morphohybridum A.Rojas. (Rojas-Alvarado 2017, 2019). Within Elaphoglossum, documented hybrids are few, but two are known to involve members of sect. Polytrichia (Lorence 1984, Rojas-Alvarado 2017). Of particular interest is E. × morphohybridum (endemic to Cocos Island), a cross between E. alvaradoanum A.Rojas and E. crinitum. Both are members of sect. Polytrichia, with the first belonging to subsect. Apoda, and the second to subsect. Polytrichia (Rojas-Alvarado 2017, Matos et al. 2021). Also of note is E. × setaceum, endemic to Mauritius, which is the only reported case of hybridization between sections in Elaphoglossum. One of its parents is E. hybridum (Bory) Brack. (sect. Polytrichia) and the other E. lanatum (Bojer ex Baker) Lorence (sect. Lepidoglossa). Hybridization may be more common and widespread in Elaphoglossum than previously detected because the morphological similarities of nearly all members of the genus makes hybrids less likely to be found. Only five species of Elaphoglossum sect. Polytrichia have been studied cytologically. Sorsa (1966) studied E. crinitum and found n = 41. That same species was studied by Walker (1966), who also found n = 41, and Walker (1985), 2n = 82. Also, Walker (1966) studied E. apodum, which was found to have n = 41, and E. denudatum, n = 41. Manton & Vida (1968) studied E. hybridum, n = 41, 2n = 82, and Walker (1985) studied E. ambiguum [as E. longifolium (Jacq.) J.Sm.], n = 41; 2n = 82. Although these species are all diploid and have the basic chromosome number of 41, it seems likely that polyploids will be found in E. sect. Polytrichia (pers. obs.). There are several reports of polyploids in other sections of the genus, including tetraploids [e.g., E. chartaceum (Baker) C.Chr., E. hirtum (Sw.) C.Chr., E. petiolatum • Phytotaxa 535 (1) © 2022 Magnolia Press MATOS ET AL. (Sw.) Urb., E. tectum (Humb. & Bonpl. ex Willd.) T.Moore], hexaploids [e.g., E. laurifolium (Thouars) T.Moore], and octoploids [e.g., E. commutatum (Mett. ex Kuhn) Alderw.] (Walker 1966, Manton & Vida 1968, and Manton unpubl. apud Manton & Vida 1968, respectively). Few molecular phylogenetic studies have included members of Elaphoglossum sect. Polytrichia. Rouhan et al. (2004), the most comprehensive phylogeny for the genus (with 120 species total), included only seven species of sect. Polytrichia. They also included E. decoratum, but this species was recovered in the Setosa clade. This result was unexpected on the basis of morphology because E. decoratum lacks hydathodes. Lóriga et al. (2014) recovered a Polytrichia clade comprising 11 species, including E. decoratum. More recently, Matos (2015) increased the sampling of sect. Polytrichia to 35 species and presented a molecular phylogeny for the group based on plastid DNA sequences (atpB-rbcL, rps4-trnS, and trnL-trnF). He also recovered sect. Polytrichia as monophyletic, with E. decoratum included. A surprising result of the Matos (2015) study was the presence in the Polytrichia clade of E. albomarginatum A.R.Sm., E. ambiguum Alston, and E. decursivum Mickel. These species are atypical in the section by lacking subulate scales. They are informally called the Decursivum clade. Their leaves appear nearly glabrous (Fig. 3A, H), and accordingly, they were previously thought to belong to Elaphoglossum sect. Elaphoglossum, whose laminae are also glabrous or beset with only scattered microscales (Mickel & Atehortúa 1980). Matos et al. (2019) studied this clade in more depth, providing a molecular phylogeny and a taxonomic revision for the group. They recognized seven species in the clade: E. albomarginatum, E. ambiguum, E. decursivum, E. lonchophyllum (Fée) T.Moore, E. miersii (Baker) C.Chr., E. papyraceum (Fée) F.B.Matos & R.C.Moran, and E. vascoae F.B.Matos & R.C.Moran. Besides lacking subulate scales, members of the Decursivum group exhibit another uncommon character in sect. Polytrichia: their vein tips are expanded laterally and often united with adjacent tips to form a submarginal vein (Matos et al. 2019). Another clade recovered by Matos (2015) and subsequently monographed is the “Apoda clade” (Matos et al. 2021). Its species are usually distinguished from others in sect. Polytrichia by the combination of brightly colored stem scales (yellowish, orangish, or reddish), evenly distributed subulate scales on laminar surfaces (Fig. 3F), and abundant tiny glandular hairs on different parts of the leaves. These characters agree with Elaphoglossum sect. Polytrichia subsect. Apoda, which was first studied by Atehortúa (1983) as the “Elaphoglossum apodum complex” (see discussion above). Matos et al. (2021) recognized 13 species in this group, including E. alvaradoanum, E. amazonicum, E. apodum, E. atehortuae F.B.Matos & R.C.Moran, E. auripilum, E. backhouseanum; E. cubense, E. latum, E. longipilosum (Atehortúa ex Mickel) A.Rojas, E. pendulum A.Rojas, E. polyblepharum Mickel, E. raywaense, and E. trichophorum (Sodiro) C.Chr. They presented descriptions, comments, synonyms (when applicable), typifications (when applicable), maps, and illustrations, as well as an identification key to all species of this group. Despite these taxonomic studies of Elaphoglosssum sect. Polytrichia, the number of species in the section remains unknown and the geographic distribution of several species poorly understood. To remedy this and as an aid to future floristicians and monographers, we provide a complete list of the currently accepted species in E. sect. Polytrichia, with places and dates of publication for the names, information on types, synonyms (when applicable), and notes on the geographic distribution of each species. For E. sect. Polytrichia as a whole, we also provide the first map of geographic distribution and a heat map showing species-richness. Methods The protologues of all names belonging to Elaphoglossum, many of which were originally published under Acrostichum, were obtained chiefly from the collections of the LuEsther T. Mertz Library of the New York Botanical Garden (http://www.nybg.org/library/). We surveyed the protologues looking for species that would fit the profile of E. sect. Polytrichia sensu Mickel & Atehortúa (1980); that is, those species described as having hydathodes absent and subulate scales present. We also searched for specimens bearing subulate scales and lacking hydathodes in the Elaphoglossum collections of the following herbaria: AAU, B, BHCB, BM, BOLV, BR, BSC, CAS, CEPEC, COL, CR, F, FLOR, FURB, GH, HAC, HAJB, HAMAB, HB, HBG, HBR, HLDG, HUA, HUCP, HUEM, HUEFS, HULE, INB, INPA, JAUM, JBSD, K, LIL, LPB, LSCR, M, MA, MBM, MEXU, MGC, MICH, MO, MSC, NY, P, PH, PI, Q, QCA, QCNE, QPLS, R, RB, S, SEL, SP, SPF, UC, ULV, UPCB, US, and YU (acronyms according to Thiers 2021). Most of the type specimens were found during these visits. We also made extensive use of web-based resources, such as Global Plants on JSTOR (https://plants.jstor.org), Reflora Programme (http://reflora.jbrj.gov.br), and Tropicos (https://www. tropicos.org/). Whenever possible, specimens were requested on loan to NY for further comparative studies. A NOMENCLATOR FOR ELAPhoGLoSSUM SECTION PoLYTrIChIA Phytotaxa 535 (1) © 2022 Magnolia Press • FIGURE 3. Some neotropical species of Elaphoglossum sect. Polytrichia. A. E. lonchophyllum, abaxial lamina base. B. E. brevipetiolatum, abaxial lamina base. C. E. glaziovii, abaxial lamina base. D. E. decoratum, abaxial lamina. E. E. mexicanum, abaxial lamina base. F. E. trichophorum, abaxial lamina base. G. E. tambillense, young leaf with hydathodes. H. E. lonchophyllum, abaxial lamina showing submarginal connecting vein. I. E. crinitum, adaxial lamina showing anastomosing veins. • Phytotaxa 535 (1) © 2022 Magnolia Press MATOS ET AL. FIGURE . Elaphoglossum crinitum (A–B) and E. decoratum (C–D). A. Habit. B. Abaxial side of sterile lamina. C. Habit. D. Abaxial side of stertile lamina. (made by Charles D. Clare on July 1973; used by Proctor 1985). A NOMENCLATOR FOR ELAPhoGLoSSUM SECTION PoLYTrIChIA Phytotaxa 535 (1) © 2022 Magnolia Press • FIGURE 5. Elaphoglossum versatile (A–D) and E. clathratum (E–J). A. Habit. B. Stem scale. C. Abaxial side of the sterile lamina. D. Scale from lamina margin. E. Habit. F. Stem scale. G. Petiole detail. H. Abaxial side of the sterile lamina. J. Scale from lamina margin. (A–D, Sodiro s.n., P; E–J, Matos 2496, QCA). 0 • Phytotaxa 535 (1) © 2022 Magnolia Press MATOS ET AL. During the last 10 years, we examined 3475 gatherings from 62 herbaria (see above). All label data were organized in a database, which was used to produce a distribution map for the section, a species richness heat map, a boxplot of elevational ranges, and the nomenclatural paragraphs. The distribution map was based on nearly all specimens examined and generated with QGIS 2.0 (QGIS Development Team 2013). Whenever possible, geographic coordinates were estimated for specimens lacking precise label data. The species richness heat map and the boxplot showing elevational ranges were generated with RStudio (Racine 2012). In the nomenclature section, we cite barcode numbers or, if these were absent on the sheets, accession numbers for the types. For some herbaria (e.g., AAU, GH, MICH) both numbers were absent, and therefore could not be cited. We also examined the scales and spores of some species using a scanning electron microscope at the Pfizer Plant Research Laboratory of the New York Botanical Garden. The spores were transferred with dissecting needles from herbarium specimens to aluminum SEM stubs where they were affixed with an asphalt adhesive. The stubs were then coated with gold-palladium alloy in a sputter-coater for 3.5 min, and structures were digitally imaged using a JEOL JSM-5410LV SEM equipped with a JEOL Orion 5410 software interface at an accelerating voltage of 10 kV. In total, we have examined the spores of about 35 species of E. sect. Polytrichia. Those of the Brazilian species were presented by Matos & Mickel (2014), those of E. mickeliorum by Matos & Moran (2017), those of the Decursivum group by Matos et al. (2019), and those of subsect. Apoda by Matos et al. (2021). The plate here presented is a selection of 12 species (including some newly imaged ones), representing the diversity of perine types within the section. Fieldwork was conducted in Brazil (2012–2019), Costa Rica (2013), Ecuador (2014), and Mexico (2014). During these trips, we observed and collected 26 of the 53 recognized species in section Polytrichia. Three other species (E. apodum, E. cubense, and E. procurrens) were photographed and collected by our collaborator Josmaily Lóriga in Cuba. Field observations have sharpened our species concepts by allowing the collection of data unavailable from herbarium specimens alone. Typification and synonymization follow the rules of the Shenzhen Code (Turland et al. 2018). Results We recognize 53 species of Elaphoglossum sect. Polytrichia (Sodiro) Christ. Acrostichum versatile is transferred to Elaphoglossum (as Elaphoglossum versatile), E. erinaceum var. occidentale is elevated to species rank (as E. occidentale); and the description of E. clathratum is expanded based on newly found fertile specimens. Line drawings are provided for E. clathratum and E. versatile (Fig. 5), as well as field photographs for E. clathratum (Fig. 6). Additionally, one epitype (E. clathratum), 13 lectotypes (A. barbatum, A. hybridum var. minor, A. mexicanum, A. nitidum, A. stramineum, E. angustioblongum, E. cinctum, E. cordifolium, E. demissum, E. erinaceum var. boliviense, E. lindbergii var. truncatum, E. reductum, E. truncatum), and two neotypes (A. hystrix, A. versatile) are designated below. No infraspecific taxa are recognized. Lastly, provided is a map of geographic distribution for the section (Fig. 7), a heat map of species richness (Fig. 8), a synopsis of species richness by country (Table 1), and a numerical list of taxa and specimens examined (Appendix 1). TABLE . Distribution by country of the species of Elaphoglossum sect. Polytrichia. Country No. spp. Species Argentina 1 hybridum Belize 3 decursivum, erinaceum, latum Bioko 1 hybridum Bolivia 12 amazonicum, barbatum, blepharoglottis, decoratum, hybridum, hystrix, kessleri, miersii, papyraceum, polyblepharum, raywaense, semisubulatum Brazil 13 amazonicum, amorimii, brevipetiolatum, decoratum, glaziovii, hybridum, miersii, papyraceum, polyblepharum, prestonii, raywaense, scolopendrifolium, ulei ......continued on the next page A NOMENCLATOR FOR ELAPhoGLoSSUM SECTION PoLYTrIChIA Phytotaxa 535 (1) © 2022 Magnolia Press • TABLE (Continued) Country No. spp. Species Cameroon Cocos Island 1 2+1 hybrid hybridum alvaradoanum, crinitum, ×morphohybridum Colombia 15 ambiguum, backhouseanum, barbatum, beitelii, crinitum, crispatum, decoratum, decursivum, erinaceum, hybridum, latum, raywaense, tambillense, vascoae, versatile Comoros 1 hybridum Costa Rica 20 albomarginatum, angustioblongum, auripilum, backhouseanum, baquianorum, barbatum, cotoi, crinitum, decoratum, decursivum, denudatum, erinaceum, gomezianum, latum, lonchophyllum, longipilosum, pendulum, reductum, silencioanum, tambillense Cuba 8 apodum, crinitum, cubense, decoratum, decursivum, denudatum, erinaceum, procurrens Democratic Republic of the Congo 1 hybridum Dominica 4 apodum, crinitum, erinaceum, tambillense Dominican Republic 9 ambiguum, apodum, crinitum, decoratum, decursivum, erinaceum, latum, lonchophyllum, tambillense Ecuador 14 atehortuae, barbatum, clathratum, crinitum, crispatum, decoratum, hybridum, latum, miersii, raywaense, tambillense, trichophorum, vascoae, versatile El Salvador 5 albomarginatum, baquianorum, erinaceum, martinezianum, mexicanum Ethiopia 1 hybridum French Guiana 1 raywaense Grenada 2 apodum, crinitum Guadeloupe 4 ambiguum, apodum, crinitum, decoratum, erinaceum Guatemala 12 albomarginatum, baquianorum, barbatum, crinitum, decoratum, decursivum, erinaceum, latum, lonchophyllum, martinezianum, mexicanum, tambillense Guyana 3 crinitum, decoratum, raywaense Haiti 8 apodum, crinitum, decoratum, decursivum, erinaceum, latum, procurrens, tambillense Honduras 6 baquianorum, decursivum, erinaceum, latum, lonchophyllum, mexicanum Jamaica 7 apodum, crinitum, cubense, decoratum, denudatum, erinaceum, tambillense Kenya 1 hybridum Madagascar 1 hybridum Malawi 1 hybridum ......continued on the next page • Phytotaxa 535 (1) © 2022 Magnolia Press MATOS ET AL. TABLE (Continued) Country No. spp. Species Martinique Mauritius 2 1+1hybrid apodum, crinitum hybridum, ×setaceum Mexico 10 albomarginatum, baquianorum, barbatum, decursivum, erinaceum, latum, lonchophyllum, mexicanum, occidentale, tambillense Montserrat 2 apodum, crinitum Mozambique 1 hybridum Nicaragua 7 backhouseanum, crinitum, decoratum, decursivum, erinaceum, latum, lonchophyllum Panama 15 albomarginatum, angustioblongum, auripilum, backhouseanum, baquianorum, crinitum, decoratum, decursivum, erinaceum, gomezianum, latum, lonchophyllum, longipilosum, silencioanum, tambillense Peru 16 amazonicum, barbatum, blepharoglottis, crispatum, decoratum, erinaceum, hybridum, hystrix, kessleri, mickeliorum, miersii, papyraceum, raywaense, tambillense, vascoae, versatile Puerto Rico 5 ambiguum, apodum, crinitum, decoratum, erinaceum Réunion 1 hybridum Rwanda 1 hybridum Saint Kitts and Nevis 2 apodum, crinitum Saint Lucia 2 apodum, crinitum Saint Vincent 2 apodum, crinitum South Africa 1 hybridum Suriname 1 raywaense Tanzania 1 hybridum Trinidad and Tobago 2 ambiguum, crinitum Tristan da Cunha 1 hybridum Uganda 1 hybridum Venezuela 11 ambiguum, barbatum, beitelii, crinitum, crispatum, decoratum, erinaceum, hybridum, ornatum, polyblepharum, raywaense Zimbabwe 1 hybridum A NOMENCLATOR FOR ELAPhoGLoSSUM SECTION PoLYTrIChIA Phytotaxa 535 (1) © 2022 Magnolia Press • 3 FIGURE . Elaphoglossum clathratum (Matos 2496, NY, P, RB, UPCB). A. Habitat near type locality. B. Habitat at type locality. C. Pendant, bluish green leaves of E. clathratum. D. Petiole with ovate scales that are patent and slightly enrolled at the base. E. Abaxial side of sterile lamina. F. Clathrate scales on lamina margin. Morphology In Elaphoglossum, subulate scales are highly distinctive and found only in sects. Polytrichia and Setosa, two clades that are sister to each other (Rouhan et al. 2004, Lóriga et al. 2014, Matos et al. 2019). Section Polytrichia is distinguished from sect. Setosa by the absence of hydathodes on the leaves of adult sporophytes—a condition interpreted as plesiomorphic (Rouhan et al. 2004, Matos et al. 2019). At first glance, these subulate scales are easily mistaken for hairs because of their slender and bristly appearance. A closer examination, however, reveals that they are actually laminate scales with strongly revolute margins. Their bases are cordate and attached at the sinuses, with lobes usually overlapped, appressed to the leaf surface, and bearing glandular projections. The attachment is slightly immersed into the leaf tissue, forming a short stalk from where a widened base supports the rest of the scale body, which is usually patent and tapered towards the apex (Bell 1951, fig. 9). The whole structure looks more or less conical • Phytotaxa 535 (1) © 2022 Magnolia Press MATOS ET AL. with a longitudinal slit (Figs. 10D, G–L). The term “bristle scales” has also been used in the literature (Bell 1951, 1955, 1956). In E. sect. Polytrichia, the subulate scales are often limited to the petioles, costae, and laminar margins, but they are also present on the laminar surfaces of E. barbatum, E. beitelii, E. crinitum, E. hybridum, E. hystrix, E. martinezianum, E. procurrens, and in 13 species of E. sect. Polytrichia subsect. Apoda (see Matos et al. 2021). Scale characters such as color, shape, size, and distribution have always been used to recognize species in sect. Polytrichia. Therefore, one of the most surprising results of the molecular phylogeny presented by Matos (2015) was the inclusion of the Decursivum Group (E. albomarginatum, E. ambiguum, and E. decursivum) in sect. Polytrichia. The best way to distinguish the Decursivum Group is its lack of costal scales, whereas scales are usually present in sect. Elaphoglossum. Also, the costal scales in sect. Elaphoglossum often have teeth, lobes, or segments, all of which are lacking on the petiole or stem scales of the Decursivum Group (Matos et al. 2019). Elaphoglossum brevipetiolatum (Fig. 3B) and E. clathratum (Figs. 5E–J, 6) also lack subulate scales, but have along their laminar margins a nearly continuous row of flat (i.e., nonsubulate), almost overlapping scales. This marginal row of flat scales is also found in E. amorimii, E. glaziovii (Fig. 3C), E. mickeliorum, E. ornatum, E. prestonii, and E. scolopendrifolium. These six species, however, have typical subulate scales on their petioles and costae. Elaphoglossum decoratum (Figs. 2E, 3D, 4C–D, 10A, H–I), E. kessleri, and E. semisubulatum (Fig. 10F) also have flat marginal scales, but their petiolar and costal scales are usually enrolled only at the base. These partially enrolled scales look broad and flat (Fig. 10H), not piliform as a typical subulate-scale (Figs. 10G, J). The opposite condition occurs in a few species, such as E. versatile (Figs. 5A–D), where the leaf scales are enrolled only at the apex. The function of these leaf scales is unknown, but it seems reasonable that they defend against herbivorous insects. They might also absorb water from the atmosphere, similar to the water-absorbing scales in bromeliads (e.g., Benzing et al. 1976) and some polypod ferns (e.g., John & Hasenstein 2017). These ideas remain to be tested. The leaves of several species of sect. Polytrichia bear minute hairs and microscales. Hairs are uniseriate and either one-celled or multicelled. They are often patent and glandular at the tip. In contrast, microscales are highly reduced scales that resemble hairs. They may be uniseriate or branched, and are usually appressed and have marginal glandular cells. In other studies, such reduced structures have also been termed “branched hairs” (Lorence & Rouhan 2004), “glandular emergences” (Bell 1955), “glandular scales” (Bell 1951), “microscales” (Matos et al. 2019, 2021), “proscales” (e.g., Matos & Mickel 2014, 2018, 2019), “squamules” (Bell 1955), and “trichomidia” (e.g., Mickel 1991, 1992, 1995, Lellinger 2002, Mickel & Smith 2004, Kessler et al. 2018). Bell (1955) reported a complete series of intermediates between glandular emergences and laminate scales in several species, suggesting that these structures are homologous. The differences between them are not always clear-cut. Both hairs and microscales, because of their small sizes, are usually visible only with a hand lens or dissecting microscope. The presence or absence of hydathodes have also been important in the subgeneric taxonomy of Elaphoglossum (e.g., Christ 1899, Mickel & Atehortúa 1980). The absence of hydathodes distinguishes E. sect. Polytrichia from E. sect. Setosa, where they are present (Fig. 1). Curiously, several species of sect. Polytrichia show functional hydathodes on the early stages of their sporophytic lives (Fig. 3G). This prompted Matos & Moran (2013) to propose that hydathodes might be ephemeral in some species of this section. Bell (1955) was probably the first to observe this phenomenon in E. apodum. Later, Atehortúa (1983) reported the same for E. crinitum and E. cubense, suggesting that “perhaps the hydathodes play an important physiological role during the early developmental stages [of sporophytes] and may have some function related to the microhabitat in which the species grow.” Since then, this has also been reported for E. clathratum (Matos & Moran 2013), E. alvaradoanum, and E. auripilum (Matos et al. 2021). The presence of hydathodes on the juvenile leaves of E. angustioblongum, E. barbatum, E. erinaceum, E. hybridum, E. occidentale, E. ulei, and E. tambillense is here reported for the first time. Venation is another important character in the taxonomy of Elaphoglossum. Almost all species of sect. Polytrichia have free veins ending near the laminar margins, and this is the condition found in most other members of the genus. Elaphoglossum crinitum is unusual by its copiously anastomosing veins (Fig. 3I, 4 A–B), a condition rare in the genus, being present in only two other species: E. crassifolium (Gaud.) Anders. & Crosby and E. pachyphyllum (Kunze) C.Chr. Several other species may have occasional anastomoses, but these are particularly common in E. hieronymi (Sodiro) C.Chr., E. macrophyllum (Mett. ex Kuhn) Christ, and E. picardae Hieron. This is a good example of homoplasy within the genus because none of these species are directly related to E. crinitum (Mickel & Atehortúa 1980, Schuettpelz & Pryer 2007). Another interesting situation occurs in the Decursivum Group (sensu Matos et al. 2019), where the vein tips are laterally expanded and often united to form a commissural vein (Fig. 3H). The spores of 12 species of Elaphoglossum sect. Polytrichia are presented in Fig. 11. These species were selected to show the morphological diversity found within the section. Spore images of about 35 other species can be seen in the papers by Moran et al. (2007), Matos & Mickel (2014), Matos & Moran (2017), Matos et al. (2019), and Matos A NOMENCLATOR FOR ELAPhoGLoSSUM SECTION PoLYTrIChIA Phytotaxa 535 (1) © 2022 Magnolia Press • 5 et al. (2021). As discussed by Moran et al. (2007), the spores of sect. Polytrichia are relatively uniform if compared to other sections. In most species examined, the perines were broadly and continuously folded, nonperforated, and nonspiny. Only E. cotoi (Fig. 11C) and E. procurrens (see Matos et al. 2021, Fig. 4K) showed narrowly folded perines; E. polyblepharum (Fig. 11J) showed small perforations; and several species showed spiny perines (see Fig. 11), as previously observed for E. prestonii by Moran et al. (2007). FIGURE . A. Worldwide distribution of Elaphoglossum sect. Polytrichia. The red dot indicates Signy Island, where spores of E. hybridum were found by Lewis Smith (2014). B. Geographic distribution of E. crinitum. C. Geographic distribution of E. decoratum. • Phytotaxa 535 (1) © 2022 Magnolia Press MATOS ET AL. Distribution and ecology Elaphoglossum sect. Polytrichia is almost entirely Neotropical, occurring from Mexico and the Antilles to Argentina and Brazil (Figs. 7A, 8). The only exception is E. hybridum (Bory) Brack., which is widespread in South America, but also occurs in Africa, Madagascar, and islands of the mid-Atlantic and Indian Oceans (Fig. 7A). Viable spores of this species have been reported from one of the South Orkney Islands of Maritime Antarctic (60°43’S, 45°38’W, see red dot in Fig. 3A) (Lewis Smith 2014), attesting to its long-distance dispersal capabilities. Its elevational range is also remarkable: 50 to 4150 m, more than any other species of Elaphoglossum of which we are aware. The occurrence in Mauritius of E. hybridum establishes the eastern range limit for the section. On this island it hybridizes with a species of sect. Lepidoglossa to form E. × setaceum Lorence (Lorence 1984). The main range of E. hybridum is in South America, suggesting at least a single long-distance dispersal from the Neotropics to other parts of the world. It probably reached the Indian Ocean region by using the volcanic islands of the mid-Atlantic Ocean and the mountains in Africa as stepping-stones, which is an idea that needs testing. In the Neotropical region, some species of sect. Polytrichia have a wide range, while others live in a very limited area. Elaphoglossum crinitum (Fig. 4A–B) and E. decoratum (Fig. 4C–D) are good examples of widely distributed species. Elaphoglossum crinitum occurs from Guatemala to Ecuador and Guyana, Cocos Island, and throughout the Antilles (Fig. 7B), whereas E. decoratum is even more widespread, occurring from Guatemala to Bolivia, Guyana, and southeastern Brazil, and in every major island of the Greater Antilles and Guadeloupe (Fig. 7C). Other widespread species, such as E. barbatum and E. erinaceum, are extremely variable morphologically and probably comprise several cryptic species. Thus, their exact geographic distribution is unknown. On the other hand, good examples of rare and narrowly distributed species occur in sect. Polytrichia. For instance, E. atehortuae, which is known from a single locality in Ecuador, E. brevipetiolatum, known from a single locality in Brazil, E. clathratum, known from two localities in Ecuador, and E. reductum A.Rojas, known from a single locality in Costa Rica. The center of diversity and endemism of sect. Polytrichia is Central America, particularly in Costa Rica and Panama, where about 20 species occur (Fig. 8). Most of these species occur on the mountains, with only E. backhouseanum and E. latum being typical of the lowlands. Elaphoglossum alvaradoanum is endemic to Cocos Island, a volcanic island ca. 550 km from the Pacific coast of Costa Rica. Elaphoglossum crinitum is the only other species of sect. Polytrichia on that island. It hybridizes with E. alvaradoanum, giving rise to the recently described E. × morphohybridum. Other extensive mountain ranges, such as the Andes and the mountains of southeastern Brazil, have also played an important role in the distribution and diversification of the group. The Andes contains ca. 20 species, but our ongoing studies suggest that more species will be found there. The mountains of eastern Brazil harbors nine species, five of which are endemic (see Matos & Mickel 2014, plus E. miersii and E. papyraceum). A well-defined clade of perhaps five species (E. amorimii, E. brevipetiolatum, E. glaziovii, E. prestonii, and E. scolopendrifolium) has diversified in these mountains (Matos 2015). The species of this clade are characterized by a nearly continuous row of planar (nonsubulate) scales on laminar margins (Matos & Mickel 2014). Only four species (Elaphoglossum amazonicum, E. decoratum, E. hybridum, and E. raywaense) occur in the intervening area between the Andes and the coastal mountains of eastern Brazil. This region is mostly covered by seasonally dry vegetation, including the Caatinga (mostly tropical thorny woodlands), the Cerrado (mostly woody savannas), and the Chaco (mostly subtropical thorny woodlands). The species occurring in this region were collected near rivers, within the more humid forests that cut through this diagonal of open formations. Amazonia harbors only two species of sect. Polytrichia: E. amazonicum and E. raywaense. A third species, E. polyblepharum, occurs disjunctly in Bolivia, Venezuela, and mountains of the Brazilian Amazon, such as Serra do Aracá and Serra Curicuriari. These mountains are outliers of the Guiana shield, a region that harbors five species of the group (E. beitelii, E. crinitum, E. crispatum, E. decoratum, and E. polyblepharum). The Greater Antilles have also played an important role in the diversification of the section. Twelve species are found there (Elaphoglossum ambiguum, E. apodum, E. crinitum, E. cubense, E. decoratum, E. decursivum, E. denudatum, E. erinaceum, E. latum, E. lonchophyllum, E. procurrens, and E. tambillense), including three endemics (E. cubense, E. denudatum, and E. procurrens). Almost all of these species occur in Hispaniola, with the exception of E. cubense (from Cuba and Jamaica) and E. denudatum (from Cuba and Jamaica, plus two recent gatherings from Costa Rica). In the Lesser Antilles, four species (E. apodum, E. crinitum, E. decoratum, and E. erinaceum) are represented in Guadeloupe, but only E. apodum and E. crinitum are more widespread and occur in nearly every major island. A NOMENCLATOR FOR ELAPhoGLoSSUM SECTION PoLYTrIChIA Phytotaxa 535 (1) © 2022 Magnolia Press • FIGURE . A. Species richness heat map of Elaphoglossum sect. Polytrichia. B. Species richness heat map of Elaphoglossum sect. Polytrichia in the Neotropics. • Phytotaxa 535 (1) © 2022 Magnolia Press MATOS ET AL. Regarding growth habit and ecological preferences, most species of Elaphoglossum sect. Polytrichia are epiphytic in wet and shaded tropical forests, with a preference for undisturbed situations. Yet some members of the E. hybridum complex (e.g., E. hybridum, E. semisubulatum, and E. tambillense) are commonly found on steep road banks with a generous amount of light. Others, like E. cotoi, are also terrestrial but grow only in shaded places inside of forests. That said, some species grow on different substrates depending on the conditions of the environment. Throughout their wide geographic distributions, E. barbatum, E. glaziovii, and E. hybridum, have been recorded as epiphytic, epipetric, and terrestrial. Some species of the Decursivum group (e.g., E. decursivum, E. lonchophyllum, E. miersii, and E. papyraceum) are noteworthy for having a strong preference for tree-fern trunks (Moran et al. 2003, Matos et al. 2019). Elaphoglossum backhouseanum, E. brevipetiolatum, and possibly E. atehortuae are epiphytic “nest-formers” (Fig. 2; Matos et al. 2018, 2021). According to Matos et al. (2018), the nest-forming habit in Elaphologlossum involves four correlated characters. First, the rhizomes are short-creeping, bearing sterile leaves close together. Second, the leaves spread outward, forming a litter-trapping “basket” or funnel-shaped rosette. Third, the lamina extends to the base of the leaf, which means that the petioles are short or absent. Fourth, a thick mat of roots, root hairs, and stem scales is often present around the stem apex and among leaf bases. This specialized morphology enables the plants to capture falling leaf litter and other debris from the canopy. As this organic matter decays, it releases nutrients to the plants (Zona & Christenhusz 2015). Additionally, the thick mass of roots, scales, and humus that is formed around the stem seems to absorb reasonable amounts of water, at least enough to keep these plants hydrated through long dry seasons. See Harrison (2018) for more information on the topic and field photos of E. backhouseanum. The elevational range of Elaphoglossum sect. Polytrichia is from nearly sea level to 4150 m, with most plants collected from 800 to 2300 m (Fig. 9). FIGURE . Boxplot depicting elevational ranges (in meters) of the species of Elaphoglossum sect. Polytrichia. The lines extending from the boxes (whiskers) indicate variability outside the upper and lower quartiles. A NOMENCLATOR FOR ELAPhoGLoSSUM SECTION PoLYTrIChIA Phytotaxa 535 (1) © 2022 Magnolia Press • FIGURE 0. Examples of scales from the laminar margins (A–F) and petioles (G–L) in Elaphoglossum sect. Polytrichia. A. E. decoratum, showing scales on both sides of laminae margin. B. E. glaziovii. C. E. ornatum. D. E. hybridum. E. E. scolopendrifolium. F. E. semisubulatum. G. E. glaziovii. H. E. decoratum. I. E. decoratum, detail of scale base. J. E. hybridum. K. E. hybridum, abaxial view showing flat and widened base with erect and enrolled scale body. L. E. hybridum, lateral view showing widened base with short stalk at attachment point. Scale bars A–L = 100 μm. 0 • Phytotaxa 535 (1) © 2022 Magnolia Press MATOS ET AL. FIGURE . Spores of some species in Elaphoglossum sect. Polytrichia. A. E. barbatum (Buchtien 5166, US). B. E. clathratum (Matos 2496, NY). C. E. cotoi (Evans 85, US). D. E. crinitum (Angulo 605, NY). E. E. decoratum (herrera 5961, MO). F. E. erinaceum (Lellinger 476, US). G. E. hybridum (Matos 2057, NY) H. E. occidentale (Pringle s.n., GH). I. E. prestonii (Brade 15514, NY). J. E. polyblepharum (Fiaschi 3250, SPF). K. E. versatile (Sodiro s.n., US). L. E. ulei (Spannagel 2, NY). Scale bars A–L = 10 μm. A NOMENCLATOR FOR ELAPhoGLoSSUM SECTION PoLYTrIChIA Phytotaxa 535 (1) © 2022 Magnolia Press • FIGURE . Elaphoglossum × setaceum, a hybrid between E. hybridum (sect. Polytrichia) and E. lanatum (sect. Lepidoglossum) from Mauritius. Label attached to the holotype sheet (Lorence 15823b, MAU-0013385), showing that a single sterile individual was found among a large population of E. lanatum, on a shady stream bank. A vigorous fertile specimen of E. hybridum was growing at the base of a tree directly overhead. The hybrid is morphologically intermediate between these two species. Nomenclatural synopsis . Elaphoglossum albomarginatum A.R.Sm., Proc. Calif. Acad. Sci., ser. 4, 40: 220. 1975. Type:—MEXICO. Chiapas: Unión Juárez, SE side of Volcán Tacaná, above Talquian, [15°05’N, 92°05’W], 2100–2200 m, 16 January 1973, D.E. Breedlove & A.r. Smith 31629 (holotype: DS-663980 in CAS-0002517!; isotypes: DS-734477 in CAS-0002518!, MEXU-00874709!, NY-00149572!). • Phytotaxa 535 (1) © 2022 Magnolia Press MATOS ET AL. Elaphoglossum pallidiforme Mickel, Novon 2: 380. 1992. Type: Panama. Panamá: 2.4 mi beyond Cerro Jefe on road to Altos de Pacora, [09°04’54”N, 79°17’27”W], 800–1000 m, 10 March 1973, T.B. Croat 22656 (holotype: NY-00149686!; isotypes: MO-3308302!, NY-00149685!, UC-1525192!). Distribution:—Mexico, Guatemala, El Salvador, Costa Rica, and Panama; (250)600–2200 m (Matos et al. 2019). Remarks:—Elaphoglossum albomarginatum is one of seven species in the Decursivum group (Matos et al. 2019). Within sect. Polytrichia, the group is atypical by lacking subulate scales and having commisural veins. Elaphoglossum albomarginatum has the relatively widest and shortest laminae of any species in the Decursivum group. Its laminae are only 2.5–4.5 times longer than broad, whereas in the other species they are 8–18 times longer than broad. Although its leaves appear glabrous, they actually bear numerous branched microscales on both sides of the lamina. The arms of these microscales are short and round, not long and hairlike as in the other species of the group. Also distinctive are the rounded laminar bases and pale whitish laminar margins. Elaphoglossum albomarginatum also resembles E. tambillense, which differs by laminae without pale margins and leaves with at least a few subulate scales. . Elaphoglossum alvaradoanum A.Rojas, Brittonia 61: 293. 2009. Type:—COSTA RICA. Puntarenas: Parque Nacional Isla del Coco, sendero a Cerro Iglesias, entre la base y la punta de Cerro Pelón, 05°31’55”N, 87°04’45”W, 530 m, 18 June 1997, A. rojas 3620 (holotype: INB-0001533964!; isotypes: K not found, MO-6843222!). Distribution:—Costa Rica, known only from Cocos Island; 0–530 m (Matos et al. 2021). Remarks:—Elaphoglossum alvaradoanum belongs to subsect. Apoda (Matos et al. 2021). The 13 species of this group are characterized by brightly colored stem scales, evenly distributed subulate scales on laminar surfaces, and tiny glandular hairs on different parts of the leaves. Elaphoglossum alvaradoanum is endemic to Cocos Island, and its first gathering is from 1898 (Pittier 12359, US). Atehortúa (1983) was the first to recognize its distinctiveness, but did not describe it as new because of the lack of fertile material. Before it was described by Rojas-Alvarado (2009), most specimens of E. alvaradoanum were identified in herbaria as E. apodum, E. auripilum var. longipilosum, or E. erinaceum. See Matos et al. (2021) for differences between these species. Rojas-Alvarado (2009) mentioned a holotype at INB and isotypes at K and MO. We found the holotype (unannotated) among specimens of E. auripilum at INB. Its label had a printed note on the upper right corner indicating that the type gathering (rojas 3620) originally comprised six duplicates (i.e., dup.=6). Unfortunately, only one of the isotypes was found (i.e., MO-6843222). A putative hybrid between E. alvaradoanum and E. crinitum was described from Cocos Island by Rojas-Alvarado (2017) as E. ×intermedium A.Rojas. This name is a later homonym of E. intermedium Brack. (syn. E. plumosum (Fée) T.Moore) from Brazil. To fix this mistake, Rojas-Alvarado (2019) published the replacement name E. × morphohybridum A.Rojas (see “Hybrids,” at the end of this paper). 3. Elaphoglossum amazonicum Atehortúa ex Mickel, Fieldiana, Bot., n.s., 27: 122. 1991. Type:—PERU. San Martín: Zepelacio, near Moyobamba, [06°02’S, 76°58’W], 1100–1200 m, October–November 1933, G. Klug 3330 (holotype: US-00048854!; isotypes: B-200072383!, F-736358!, GH-00967239!, K-000501544!, MO-1081993!, NY-00149574!, S-13-13365!, USM-000475 [image!]). Distribution:—Peru, Bolivia, and W Brazil; 490–1200 m (Matos et al. 2021). Remarks:—Elaphoglossum amazonicum belongs to subsect. Apoda (Matos et al. 2021). It was first recognized by Underwood, who annotated a gathering from Bolivia (Williams 1027, GH, NY) as E. williamsii Underw., but never published the name. When Atehortúa (1983) studied this group, the epithet “williamsii” was already unavailable because of E. williamsii Vareschi [=E. nigrescens (Hook.) T.Moore ex Diels], so she proposed E. amazonicum Atehortúa (nom. inval.). Mickel (1991) later validated this name because Atehortúa’s doctoral thesis does not constitute an effective publication (Turland et al. 2018, Art. 30.9). . Elaphoglossum ambiguum Alston, J. Wash. Acad. Sci. 48: 234. 1958. Acrostichum ambiguum Mett. ex Christ, Neue Denkschr. Allg. Schweiz. Ges. Gesammten Naturwiss. 36: 60. 1899, nom. illeg. Type:—VENEZUELA. [Aragua]: “Prope coloniam Tovar,” [10°24’20”N, 67°17’22”W], 1950 m, [without day or month] 1854–1855, A. Fendler 283 (lectotype, designated by Kessler et al. 2018: B-200069755!; isolectotypes: BR-0000006965242!, K-000501546!, MO-1867368!, US-00067315!, YU-000646!, YU-000647!). A NOMENCLATOR FOR ELAPhoGLoSSUM SECTION PoLYTrIChIA Phytotaxa 535 (1) © 2022 Magnolia Press • 3 Distribution:—Dominican Republic, Guadeloupe, Trinidad, Venezuela, and Colombia; 610–2300 m (Matos et al. 2019). Remarks:—Elaphoglossum ambiguum belongs to the Decursivum Group (Matos et al. 2019). The seven species of this group are unusual in sect. Polytrichia for lacking subulate scales and for having commisural veins. Elaphoglossum ambiguum most resembles E. vascoae, from which it differs by the petiole scales that are firm, lanceolate with acuminate apices, and composed of elongate cells. In contrast, those of E. vascoae are weak and flaccid, of rather indistinct form but with obtuse apices, and composed of isodiametric cells. Additionally, the petiolar scales of E. ambiguum are usually spreading, whereas those of E. vascoae are tightly appressed and wrapped around the petioles. The cell shape in the petiole scales of the Decursivum group can be seen in fig. 5 of Matos et al. (2019). However, the caption of that figure is actually wrong: E. ambiguum should be 5A, E. decursivum 5B, E. lonchophyllum 5C, E. miersii 5D, E. papyraceum 5E, and E. vascoae 5F. Determining the correct name for this species has had a tortuous history fully detailed by Kessler et al. (2018). The treatment of this species for Bolivia (Kessler et al. 2018), however, was based on specimens that we now consider to be an undescribed species of E. sect. Elaphoglossum sensu Rouhan et al. (2004). 5. Elaphoglossum amorimii F.B.Matos & Mickel, Brittonia 66: 377. 2014. Type:—BRAZIL. Bahia: Camacan, Fazenda Serra Bonita, 9.7 km W de Camacan na estrada para Jacarecí, daí 6 km SW na estrada para RPPN Serra Bonita e torre da Embratel, 15°23’30”S, 39°33’55”W, 835 m, 9 July 2005, F.B. Matos, A. Amorim, J. Lima, A. Lobão & S. Sant’Ana 620 (holotype: CEPEC-105144; isotype: UPCB-52941). Distribution:—E Brazil, endemic to Bahia and Espírito Santo; 530–1000 m. Remarks:—Elaphoglossum amorimii most resembles E. glaziovii and E. prestonii, which are also endemic to eastern Brazil. They all have petiolar and costal scales that are subulate, and lamina margin scales that are flat. See Matos & Mickel (2014) for comments on how to distinguish these species. . Elaphoglossum angustioblongum A.Rojas, Revista Biol. Trop. 51: 34. 2003, as “angustiobongum” in fig. 1. Type:—PANAMA. Chiriquí: Las Cumbres, hogback ridge N of Quebrada Iglesia, near town of Cerro Punta, [8°51’N, 82°34’W], 22 July 1971, T.B. Croat & D. Porter 16078 (holotype: MO, currently lost). Lectotype:—PANAMA. Chiriquí: Distrito Bugaba, Cerro Punta, 8°52’N, 82°33’W, 2200 m, 23 January 1984, h. van der Werff & J. herrera 6271 (lectotype, designated here: MO-3228579!). (Fig. 2A) Distribution:—Costa Rica and Panama; (400–)1100–2500 m. Remarks:—The distinctness of this taxon was first recognized by H. Christ, who annotated a gathering from Navarro, Costa Rica (Wercklé s.n., P-01466158) as “Elaphoglossum navarrense sp. nov.”, but never published the name. We were not able to locate the holotype of E. angustioblongum at MO, even though we have searched that herbarium several times. To fix the application of this name, we designate one of the paratypes as lectotype. The specimen selected is sterile, but otherwise complete. It was collected near the holotype locality and is a good match for the description. Two other paratypes were found at MO (Davidse 24334, Werff & herrera 6303) and can be used to understand the concept of this species. The type of E. demissum Christ, which is not a subulate-scaled species, was mounted on a mixed sheet containing three juvenile specimens of E. angustioblongum. See the excluded species section for additional comments. . Elaphoglossum apodum (Kaulf.) Schott ex J.Sm., J. Bot. (Hooker) 4: 148. 1841. Acrostichum apodum Kaulf., Enum. Filic.: 59. 1824. Type:—MONTSERRAT. [Without precise locality], s.d., J. ryan s.n. (lectotype, designated by Matos et al. 2021: BM-001073357!). Epitype:—SAINT VINCENT. [Without precise locality], [without day or month] 1827, rev. L. Guilding s.n. (epitype, designated by Matos et al. 2021: E-00690737!; isoepitypes: K-000501579!, K000501580!, K-000501581!). Acrostichum platyneuron Fée, Mém. Foug., 2. Hist. Acrostich.: 43. 1845, nom. illeg., non A. platyneuron L., Sp. Pl.: 1069–1070. 1753, as “platyneuros”. Elaphoglossum platyneuron T.Moore, Index Fil.: 13. 1857. Type:—CUBA. Santiago de Cuba: La Guinea, le Gato et la Pelada, [20°24’43”N, 75°31’43”W], August 1844, J. Linden 2156 (lectotype, designated by Matos et al. 2021: P-01818748!; isolectotypes: BR-0000013075033!, GENT-10170304! [image!], NY-00127326!, P-00249917!, P-01818747!), as “J. Linden 2056.” • Phytotaxa 535 (1) © 2022 Magnolia Press MATOS ET AL. Distribution:—Cuba, Jamaica, Haiti, Dominican Republic, Puerto Rico, Saint Kitts and Nevis, Montserrat, Guadeloupe, Dominica, Martinique, Saint Lucia, Saint Vincent, and Grenada; 50–1200 m (Matos et al. 2021). Remarks:—Elaphoglossum apodum belongs to subsect. Apoda (Matos et al. 2021). The name has been widely applied in the neotropics, but E. apodum is endemic to the Greater and Lesser Antilles. Nearly all specimens annotated with this name from Central and South America refer to E. latum or E. raywaense (Matos et al. 2021). The holotype of Acrostichum apodum was probably lost in a shipment from Halle to Copenhagen (Matos et al. 2021). A duplicate of the holotype was found at BM (ryan s.n., BM-001073357), and this specimen was later designated as lectotype of A. apodum by Matos et al. (2021). These authors also designated an epitype (Guilding s.n., E-00690737), particularly because the lectotype lacked a stem, which provides critical characters for the precise application of the name. . Elaphoglossum atehortuae F.B.Matos & R.C.Moran, Syst. Bot. 46: 779. 2021. Type:—ECUADOR. MoronaSantiago: Cantón Huamboya, Shankarimi, 01°58’S, 77°49’W, 850 m, 2 August 1993, W.A. Palacios 10963 (holotype: MO-05054610!; isotype: QCNE, not found). Distribution:—Amazonian Ecuador, known only from the holotype; 850 m (Matos et al. 2021). Remarks:—Elaphoglossum atehortuae belongs to subsect. Apoda (Matos et al. 2021). The species was described based on a single sterile specimen from Ecuador. According to the collector of the type (W. Palacios, pers. comm.), there should be an isotype at QCNE, but this material is currently lost. The most similar species is E. backhouseanum, which occurs from Nicaragua to NW Colombia (see Matos et al. 2021 for a comparison). . Elaphoglossum auripilum Christ, Bull. Herb. Boissier, sér. 2, 5(1): 8. 1905. Type:—COSTA RICA. [Without precise locality], [without day or month] 1903, C. Wercklé s.n. (lectotype, first-step designated by Mickel, Fl. Mesoamer. 1: 258. 1995, second step designated by Matos et al. 2021: P-00249661!; isolectotype: P-00249662!). Distribution:—Costa Rica and Panama; 1000–2400(–2755) m. (Matos et al. 2021). Remarks:—Elaphoglossum auripilum is one of the most variable species of subsect. Apoda (Matos et al. 2021). Mickel (1992) described E. auripilum var. longipilosum Atehortúa’s ex Mickel based on specimens with “more abundant, longer scales on petiole (3–6 mm vs. 1–3 mm long) and lamina (2–3 mm vs. 1.5–2 mm long)”. RojasAlvarado (2009) elevated this variety to species rank, and the name E. longipilosum A.Rojas was recognized by Matos et al. (2021). 0. Elaphoglossum backhouseanum T.Moore, Gard. Chron., n.s., 17: 672. 1882, as “Backhousianum”. Type:— [Locality and collector unknown] (holotype: K-000590606!). Elaphoglossum gerardianum L.D.Gómez, Phytologia 60: 74. 1986. Type:—COSTA RICA. Limón: [Sukut], camino entre la finca de don Calixto Kiamble y el antiguo camino a Katsi, subiendo hasta el Cerro Kikírchabeta, [09°25’N, 82°58’W], 500 m, 23 October 1985, L.D. Gómez, G. herrera & D. Masterson 23784 (holotype: CR-115229!; isotypes: CR-124413!, CR-222666!). (Fig. 2B) Distribution:—Nicaragua, Costa Rica, Panama, and NW Colombia; 35–850 m (Matos et al. 2021). Remarks:—Elaphoglossum backhouseanum belongs to subsect. Apoda and is unusual in that group by lacking subulate scales on both laminar surfaces (Matos et al. 2021). It is one of the few nest-former species in the genus, having large oblanceolate leaves that are subsessile and ascendant, forming a basket that captures litter from the canopy (Harrison 2018, Matos et al. 2018, 2021). The holotype of E. backhouseanum was said to be from Mexico (Moore 1882), but this species does not occur in that country (Mickel & Smith 2004). Mickel (1995) suggested that the type came from Panama, but the earliest collections of E. backhouseanum are from Nicaragua (March 1893, Shimek s.n., ISC) and Costa Rica (April 1895, Tonduz 9449, US), with only recent collections from Colombia and Panama (Matos et al. 2021). . Elaphoglossum baquianorum A.Rojas, Revista Biol. Trop. 51: 35. 2003. Type:—COSTA RICA. Puntarenas: Cordillera de Talamanca, Parque Internacional La Amistad, sendero a Cerro Kámuk, entre Cerro Kasir y Cerro Nai, 9°11’50’’N, 83°03’30’’W, 2900–3100 m, 8 November 1996, A. rojas 3208 (holotype: INB-0001520922!; isotypes: CR-248097!, MEXU?, MO-6745098 [image!], NY?, UC?, US?). A NOMENCLATOR FOR ELAPhoGLoSSUM SECTION PoLYTrIChIA Phytotaxa 535 (1) © 2022 Magnolia Press • 5 Distribution:—Mexico, Guatemala, Honduras, El Salvador, Costa Rica, and Panama; 2000–3300 m. Remarks:—The differences between Elaphoglossum baquianorum and E. tambillense are few and subtle (see Rojas-Alvarado 2003), but a recent phylogenetic study based on plastid DNA data (Matos 2015) suggests that they are not closely related. Rojas-Alvarado (2003) mentioned isotypes at CR, MEXU, MO, NY, UC, and US, but we only found them at CR and MO. . Elaphoglossum barbatum (H.Karst.) Hieron., Bot. Jahrb. Syst. 34: 553. 1904. Acrostichum barbatum H.Karst., Fl. Columb. 2(5): 155. 1869, non Mettenius (1864), nom. nud. Type:—COLOMBIA. Bogota: 2200–2800 m, s.d., G.K.W.h. Karsten s.n. (lectotype, designated here: LE-00008851!; isolectotypes: B-200069894!, LE-00008850!, P-00249835!). (Fig. 11A) Elaphoglossum erinaceum var. boliviense Rosenst., Repert. Spec. Nov. Regni Veg. 12: 476. 1913, as “boliviensis”. Type:—BOLIVIA. La Paz: Nor Yungas, Polo-Polo prope Coroico, [16°11’S, 67°44’W], 900–1100 m, October–November 1912, o. Buchtien 3470 (lectotype, designated here: S-05-9346!; isolectotypes: F-515211!, NY-00149616!, US-00067340!). Elaphoglossum cordifolium Rosenst., Repert. Spec. Nov. Regni Veg. 25: 62. 1928. Type:—BOLIVIA. La Paz: Larecaja, Hacienda Casana sobre el camino a Tipuani, [15°38’S, 68°17’W], 1400 m, 27 January 1923, o. Buchtien 7040 (lectotype, designated here: UC478078!; isolectotypes: S-R-1844!, US-00067334!). Elaphoglossum truncatum Rosenst., Repert. Spec. Nov. Regni Veg. 25: 63. 1928. Elaphoglossum lindbergii var. truncatum Rosenst., Repert. Spec. Nov. Regni Veg. 25: 63. 1928. Type:—BOLIVIA. La Paz: Larecaja, Hacienda Simaco sobre el camino a Tipuani, [15°37’S, 68°04’W], 1400 m, February 1920, o. Buchtien 5166 (lectotype, designated here: UC-478081!; isolectotypes: NY00149657!, MO-5384661!, US-00067355!, US-00955782!). Other syntypes:—BOLIVIA. La Paz: Larecaja, Hacienda Casana, sobre el camino Tipuani, [15°38’S, 68°17’W], 1400 m, 27 January 1923, o. Buchtien 7042 (UC-478079!, US-1499170!). Distribution:—Mexico, Guatemala, Costa Rica, Colombia, Venezuela, Ecuador, Peru, and Bolivia; 700–3300 m. Remarks:—According to Tryon (1963), LE has Karsten’s most complete set of ferns from South America. There are two sheets of Acrostichum barbatum at LE, and the label data on both correspond to the protologue. We designate LE00008851 as the lectotype because it is the only sheet with a stem. Also, this sheet has an authentic label embossed with “H. Karsten” and the specimen is a good match for the illustration depicted in Florae Columbiae (Karsten 1869: tab. 181). One of the three fertile leaves of this specimen is atypical in having only its upper third covered by sporangia. Lindig 111 (B) is not a type, although it has been cited as such by a few authors (e.g., Mickel 1991). This confusion was probably caused by Mettenius (1864), who cited this collection under A. barbatum five years before the name was validly published. The types of Elaphoglossum barbatum, E. cordifolium, and E. truncatum are quite distinct. The main difference between them is the shape of the lamina base, which varies from cuneate to strongly cordate. However, we do not recognize them as distinct because there is a complete series of intermediate specimens between each of these types. The presence of subulate scales on the laminar surfaces, which is often used to distinguish E. barbatum from its closely related species, is also variable. However, this character is difficult to assess because these scales are often caducous or easily abraded. Further studies at a populational level may show that E. barbatum comprises more than one species. rojas 2856 (INB) and other collections that have been annotated as “E. hyposquamatum A.Rojas” represent this species. The name was never effectively published. Buchtien 5166 and 7042 are syntypes of two legitimate names that were simultaneously published (i.e., Elaphoglossum lindbergii var. truncatum and E. truncatum). Elaphoglossum cordifolium and E. truncatum have equal priority (Turland et al. 2018, Art. 11.5), and this must be considered if someone decides that populations from Bolivia are not the same as the earlier E. barbatum from Colombia. 3. Elaphoglossum beitelii (Mickel) A.Rojas, Revista Biol. Trop. 50: 984. 2002. Elaphoglossum crispatum Mickel var. beitelii Mickel, Brittonia 39: 316. 1987. Type:—VENEZUELA. Amazonas: Depto. Río Negro, Cerro de la Neblina, 5.1 km NE Pico Phelps, 21.5 km E Neblina Base Camp, 00°50’40”N, 65°58’10”W, 1850 m, 3 February 1985, J.M. Beitel 85133 (holotype: NY-00149604!; isotypes: MO-4251772!, UC-1551700!). Distribution:—Colombia and Venezuela; 1920–2500 m. Remarks:—Elaphoglossum beitelii is known only from two gatherings. It is characterized by dark (i.e., castaneous to black) and crispate stem scales, narrowly oblong laminae with cuneate bases, acuminate apices, and subulate scales persistent on laminar surfaces. • Phytotaxa 535 (1) © 2022 Magnolia Press MATOS ET AL. . Elaphoglossum blepharoglottis Mickel, Fieldiana, Bot., n.s., 27: 126. 1991. Type:—PERU. Huánuco: Muña, [09°40’S, 75°49’W], 2100 m, 23 May–4 June 1923, G.S. Bryan 534 (F-1829444!). Distribution:—Peru and Bolivia; 1250–2100 m. Remarks:—Previously, this species was known only from the holotype, but two more recent collections are here reported (Kessler 13359 from Bolivia, and Mellado 2312 from Peru, both at NY). Elaphoglossum blepharoglottis is characterized by its large, narrowly oblanceolate leaves with attenuate laminar bases and subulate scales mostly limited to petioles, costae, and laminar margins. These scales are dark brown to black, strongly subulate, and dense. The ones on the petioles and costae are patent, but the ones along the laminar margins are smaller and strongly ascending. The specimen collected by Mellado 2312 (NY) differs from the holotype in having numerous subulate scales on the abaxial laminar surfaces. It superficially resembles E. trichophorum from Ecuador, but has darker stem scales. 5. Elaphoglossum brevipetiolatum F.B.Matos & Mickel, Brittonia 66: 384. 2014. Type:—BRAZIL. Espírito Santo: Santa Teresa, Nova Lombardia, Reserva Biológica Augusto Ruschi, [19°54’58”S, 40°32’31”W], 900 m, 1 October 2002, r.r. Vervloet, E. Bausen, W. Pizziolo & J. rosini 1112 (holotype: MBML-17941!; isotype: RB-429725!). (Fig. 3B) Distribution:—SE Brazil, known only from Espírito Santo; 800–900 m. Remarks:—This narrow endemic species from the mountains of southeastern Brazil is one of the few members of section Polytrichia without subulate scales. It resembles Elaphoglossum glaziovii, but its petioles vary from nearly absent to 1.5 cm long (vs. more than 2 cm long), and its petiolar and costal scales are flat (vs. subulate) (Matos & Mickel 2014). . Elaphoglossum clathratum F.B.Matos & R.C.Moran, Nordic J. Bot. 31: 442. 2013. Type:—ECUADOR. Tungurahua: road north of El Topo, Km 9, at suspension bridge over Río Topo, 1°21’S, 78°12’W, 1550–1600 m, 30 March 1998, B. Øllgaard & h. Navarrete 3000 (holotype: QCA-111250!; isotype: AAU-sheet without number!). (Figs. 5E–J, 6A–F, 11B) Distribution:—Ecuador, eastern side of the Andes; 1500–2100 m. Remarks:—Elaphoglossum clathratum is currently known from three gatherings made on the eastern side of the Andes in Ecuador. Its description was based on two of the gatherings (Øllgaard 3000 and Moran 7440), and both were incomplete, without stems and fertile leaves. In August 2014, one of us (FM) visited the type locality (Fig. 6B) and found fertile specimens (Matos 2496, QCA, NY, RB, UPCB). The fertile leaves are much shorter than the sterile ones, about 1/3 the length (Fig. 5E), and the spores have perispores with broad continuous folds and rugose surfaces (Fig. 11B). Elaphoglossum clathratum is one of the few species of E. sect. Polytrichia without typical subulate scales. It is similar to E. decoratum in having a nearly continuous band of ovate scales on laminar margins (Fig. 6E), and petiole scales that are patent and slightly enrolled only at the base (Fig. 6D). It differs from that species, however, by costal and petiolar scales brown (vs. orange), sparse (vs. dense), and caducous (vs. persistent) (Figs. 6D–E). . Elaphoglossum cotoi A.Rojas, Revista Biol. Trop. 51: 37. 2003. Type:—COSTA RICA. San José: Cantón de Dota, Cordillera de Talamanca, San Gerardo, 09°32’50”N, 83°49’50”W, 2300–2400 m, 17 July 1996, A. rojas & M. Coto 2790 (holotype: INB-0001514452!; isotypes: CR-0246522!, MO?, NY?, UC?). (Figs. 2C, 11C) Distribution:—Costa Rica (also reported from Panama by Rojas, 2003, but see comments below); 1900–3200 m. Remarks:—Three of the isotypes were not found, but it is possible that they are still being processed by their respective institutions. Rojas-Alvarado (2003) cited one collection from Panama (rojas 3286, INB, NY), but this gathering is from Costa Rica and represents E. auripilum. Some specimens of E. cotoi resemble E. erinaceum, but these two species differ in habitat. Elaphoglossum cotoi is terrestrial (vs. epiphytic) and has a crown or semi-crown of arching sterile leaves surrounding the stem apex. The fertile leaves are usually erect at the center of the crown, with petioles ca. twice as long as those of the sterile leaves (vs. shorter or about the same size as those of sterile leaves in E. erinaceum). Also, E. cotoi has lanceolate (vs. broadly oblong or elliptic) sterile laminae, sterile laminar bases rounded to decurrent from abruptly contracted laminae (vs. cuneate), and nearly glabrous (vs. evenly scaly) costae. . Elaphoglossum crinitum (L.) Christ, Neue Denkschr. Allg. Schweiz. Ges. Gesammten Naturwiss. 36: 102. 1899. Acrostichum crinitum L., Sp. Pl. 2: 1068. 1753. olfersia crinita (L.) C. Presl, Tent. Pterid.: 234. 1836. hymenodium A NOMENCLATOR FOR ELAPhoGLoSSUM SECTION PoLYTrIChIA Phytotaxa 535 (1) © 2022 Magnolia Press • crinitum (L.) Fée, Mém. Foug. 2: 90. 1845. Dictyoglossum crinitum (L.) J.Sm., Bot. Mag. 72: 18. 1846. Anetium crinitum (L.) C.Presl, Abh. Königl. Böhm. Ges. Wiss., ser. 5, 6: 536. 1851. Chrysodium crinitum (L.) Mett., Fil. Hort. Bot. Lips. 1: 21. 1856. Type:—(holotype: Petiver, Pteri-graphia Amer.: 145, tab. 13, fig. 14. 1712). Epitype:— MARTINIQUE. Morne de la Calebasse, s.d., C. Plumier s.n. [herb. Tournefortianum 5372] (epitype, designated by Cremers & Aupic 2008: P-00307150!). (Figs. 2D, 3I, 4A–B, 11D) Distribution:—Guatemala, Nicaragua, Costa Rica, Panama, Cuba, Jamaica, Haiti, Dominican Republic, Puerto Rico, Saint Kitts and Nevis, Montserrat, Guadeloupe, Dominica, Martinique, Saint Lucia, Saint Vincent, Grenada, Trinidad and Tobago, Venezuela, Guyana, Colombia, Ecuador; (60–)200–1500(–1800) m. (Fig. 7B). Remarks:—The description of this species was based exclusively on an illustration published by Petiver (1712). Proctor (1977) indicated this figure as “type” and noted that it was copied from Plumier (1705, tab. 125), which represents a plant from Martinique. Jarvis (2007) noted that no specimen of A. crinitum exists at LINN and interpreted Proctor’s citation of the Petiver plate as a lectotypification. However, this illustration is not a lectotype; it is the holotype because it was the only element upon which the validating description was based (Matos et al. 2015). Cremers & Aupic (2008) designated an epitype, which we consider helpful because the holotype does not show the reticulate venation characteristic of this species. There are other species of Elaphoglossum with anastomosing veins (e.g., Elaphoglossum crassifolium, E. hieronymi, E. hymenodiastrum, and E. pachyphyllum), but none of these have subulate scales. A putative hybrid between E. crinitum and E. alvaradoanum was recently described from Cocos Island, Costa Rica (see E. × morphohybridum A.Rojas at the end of this paper, under “Hybrids”). . Elaphoglossum crispatum Mickel, Brittonia 39: 316. 1987. Type:—VENEZUELA. Mérida: Distrito Rangel, roadside between Santo Domingo and La Soledad, 8°52’N, 71°34’W, 1500 m, 5 August 1983, h.h. van der Werff & r. ortíz 5918 (holotype: MO-3123077!; isotype: MO-3232627!). Distribution:—Venezuela, Colombia, Ecuador, and Peru; 800–2050 m. Remarks:—Elaphoglossum crispatum more closely resembles E. angustioblongum in its castaneous to black, lustrous stem scales, subcoriaceous to coriaceous laminae, and dark subulate scales mostly limited to petioles and laminar margins. It differs from E. angustioblongum, however, by strongly crispate (vs. nearly stiff) stem scales. 0. Elaphoglossum cubense (Mett. ex Kuhn) C.Chr., Index Filic.: 305. 1905. Acrostichum cubense Mett. ex Kuhn, Linnaea 36: 46. 1869. Type:—CUBA. [Guantánamo]: “in Cuba Orientali”, [20°52’N, 76°54’W], September 1859– January 1860, C. Wright 1040 (lectotype, designated by Matos et al. 2021: B-200070282!; isolectotypes: BM000769774!, E-00106047 [image!], GH-00020349!, GOET-010309 [image!], K-000590615!, LE-00008853 [image!], MA-696216 [image!], MO-1867291!, P-01464564!, PH-611349!, US-00067271!, YU-000650!). Distribution:—Cuba and Jamaica; 350–1790 m (Matos et al. 2021). Remarks:—Elaphoglossum cubense belongs to subsect. Apoda (Matos et al. 2021). Both Atehortúa (1983) and Proctor (1985) cited the B specimen as holotype. However, it is impossible to establish if that was the only element upon which the description was based, and thus all duplicates of this gathering should be considered syntypes. The designations by those authors were not inferential lectotypifications because Atehortúa (1983) does not constitute effective publication (Turland et al. 2018, Art. 7.10) and Proctor (1985) was doubtful regarding this type (i.e., his indication of a holotype was followed by a question mark) (Turland et al. 2018, Art. 7.11). Matos et al. (2021) were the first to designate the B specimen as lectotype. . Elaphoglossum decoratum (Kunze) T.Moore, Index Fil.: 8. 1857. Acrostichum decoratum Kunze, Linnaea 9: 25. 1834. Type:—PERU. Huánuco: Pampayaco, [9°33’S, 75°52’W], July 1829, E.F. Poeppig s.n. (Diar. 1134) (lectotype, designated by Mickel 1991: MO-1867358!). (Figs. 2E, 3D, 4C–D, 10A, H–I, 11E) Distribution:—Cuba, Jamaica, Haiti, Dominican Republic, Puerto Rico, Guadeloupe, Guatemala, Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Guyana, Ecuador, Peru, Bolivia, and Brazil; 300–2600 m. (Fig. 7C) Remarks:—Elaphoglossum decoratum is one of the most distinctive species of the genus, as evidenced by the fact that Mickel & Atehortúa (1980) classified it in its own section (i.e., sect. Decorata Mickel & Atehortúa). It was not until 2004, with the publication of the first DNA-based phylogenies of the genus (Rouhan et al. 2004, Skog et al. 2004), that the close relationship of E. decoratum with the subulate-scaled species was discovered. This was • Phytotaxa 535 (1) © 2022 Magnolia Press MATOS ET AL. unexpected on the basis of morphology because the scales of E. decoratum are broadly ovate and appear flat, unlike the typical subulate scales of the other species (which are hairlike and strongly enrolled lengthwise). A reexamination of these scales, however, showed that the petiolar and costal scales of E. decoratum are actually enrolled at the base (Figs. 10H–I); in fact, this character was well illustrated by Christ (1899, fig. 9). Elaphoglossum decoratum is further characterized by a nearly continuous band of orbiculate, planar, and overlapping scales along the margins of sterile laminae. The specimen W-0052454, which was annotated as “typus probabiliter” by L. Pignotti (W) in 2014, is not a type. Although it was also collected by Poeppig in Peru, the information on its original label does not agree with the protologue. It was collected in Cuchero (vs. Pampayaco), November 1829 (vs. July 1829). . Elaphoglossum decursivum Mickel, Brittonia 32: 334. 1980. Type:—MEXICO. Oaxaca: Distrito Ixtlán, 29 km S of Valle Nacional, 80 km N of Ixtlán de Juárez, trail E of Rte. 175 at Campamento Vista Hermosa toward Ladú, 1 hr. hike down to Río de la Trucha, [17°31’N, 96°25’W], 457–610 m, 13 August 1971, J.T. Mickel 6417 (holotype: NY00149607!; isotypes: MEXU-00866297!, MICH-1191296!, NY-00260467!, UC-1734404!). Distribution:—Cuba, Dominican Republic, Haiti, Mexico, Belize, Guatemala, Honduras, Nicaragua, Costa Rica, Panama, and Colombia; 450–2200 m (Sánchez 2017, Matos et al. 2019). Remarks:—Elaphoglossum decursivum belongs to the Decursivum group (Matos et al. 2019). It differs from the other six species of this group by basally long-decurrent laminae and often short petioles compared to the lamina length. Also, the cells of the petiole scales in E. decursivum are elongate and sinuate (Matos et al. 2019, fig. 5B), unlike the other species in the clade. Choo et al. (2014) studied the gametophytes and young sporophytes of this species and discovered that its earliest sporeling leaves were covered with minute papillate-glandular hairs. These hairs would decrease in abundance with each successive new leaf, becoming completely absent by the time that the young sporophytes produced leaves of about 2 cm long. Being unaware of the phylogenetic position of E. decursivum, Choo et al. (2014) reported their finding as the first observation of hairs in the glabrous sect. Elaphoglossum. Since then, molecular phylogenetic studies have shown that E. decursivum belongs to sect. Polytrichia (Matos 2015, Matos et al. 2019). This was unexpected on the basis of morphology because of the lack of subulate scales, but the presence of glandular hairs supports this relationship because such hairs are almost exclusive to the subulate-scaled species in Elaphoglossum (see Mickel & Atehortúa 1980). Mickel (1980) stated that the holotype of Elaphoglossum decursivum is at NY, with isotypes at ENCB, NY, UC, and US. We did not find the specimens at ENCB and US, but there are isotypes at MEXU and MICH. The original labels of the two NY sheets were clearly indicated by the author as different categories of types (i.e., holotype and isotype), so there is no need for a lectotypification. 3. Elaphoglossum denudatum (Jenman) Maxon ex C.V.Morton, Amer. Fern J. 52: 149. 1962. Acrostichum hybridum var. denudatum Jenman, Bull. Dept. Bot. Jamaica, n.s., 5: 89. 1898. Type:—JAMAICA. [Precise locality unknown]: [without day or month] 1875, G.S. Jenman 7 (lectotype, designated by Proctor 1985: K-000590614!; isolectotypes: NY-02008138!, NY-02008140!, NY-02008151!, NY-02008153!). Distribution:—Costa Rica, Cuba, and Jamaica. Probably more widespread, but overlooked; 300–1200 m. Remarks:—Jenman (1898) did not cite specimens in the protologue. Consequently, Morton (1962) proposed as the lectotype r.V. Sherring s.n. (BM, duplicate at US). There is nothing indicating that Jenman used this collection for the description, so it probably does not constitute original material. Following Article 9.12 of the Shenzhen Code (Turland et al. 2018), we have adopted the lectotype designated by Proctor (1985) (K000590614!), which was collected by G.S. Jenman in Jamaica and bears a label with his handwritten notes. This herbarium sheet comprises two different gatherings. The specimen on top (K000590613) was collected by W. Jameson in Ecuador and must be disregarded as type. NY-02008138 is a small fragment of the lectotype. Three other sheets (NY-02008140, 02008151, 02008153) are almost certainly part of the same gathering and were considered isolectotypes. They have a printed label reading “Jenman’s Jamaica Ferns, &c., Type Specimen–Collected 1874–1879” and were marked in Jenman’s handwriting as “Acrostichum denudatum Jenman”. The only problem is that they were not marked as number 7. Rojas-Alvarado (1996b) was the first to cite this species for Costa Rica. We are here confirming this report based on morphology and also on the results of a recent molecular phylogenetic study (Matos 2015), in which a specimen from Costa Rica (Matos 2082, NY) was recovered as sister to typical E. denudatum from Jamaica (Anderson 3304, US). A NOMENCLATOR FOR ELAPhoGLoSSUM SECTION PoLYTrIChIA Phytotaxa 535 (1) © 2022 Magnolia Press • . Elaphoglossum erinaceum (Fée) T.Moore, Index Fil.: 9. 1857. Acrostichum erinaceum Fée, Mém. Foug. 2: 41. 1845. Type:—GUADELOUPE. Rivière de Saint-Louis, s.d., de Thiouville s.n. (lectotype, designated by Proctor 1977: P-01818776!). (Fig. 2F, 11F) Acrostichum crinitum M.Martens & Galeotti, Mém. Foug. Mexique 25. 1842, nom. illeg., non A. crinitum L., Sp. Pl. 2: 1068. 1753. Acrostichum blepharodes Fée, Mém. Foug. 2: 48. 1845, nom. nov. Elaphoglossum blepharodes (Fée) T.Moore, Index Fil.: 7. 1857. Type:—MEXICO. Veracruz: prope Xalapa, [19°32’24”N, 96°55’39”W], 2000–3000 m, June–October 1840, h. Galeotti 6297 (lectotype, designated by Mickel & Beitel 1988: BR-0000006964139!; isolectotypes: P-00249704!, P-00249705!). Acrostichum nitidum Liebm., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 5, 1: 168. 1849. Elaphoglossum fimbriatum T. Moore, Index Fil.: 356. 1862, nom. nov., non Elaphoglossum nitidum Brack., U.S. Expl. Exped., Filic. 16: 70. 1854. Type:—MEXICO. Puebla: Barranca de Chinautla, [19°50’N, 97°23’W], 2133 m, [without day or month] 1841–1843, F.M. Liebmann s.n. (lectotype, first step designated by Mickel & Beitel 1988, second step designated here: C no. LNR-17750!; isolectotypes: C no. LNR-17751!, P-00249888!, US-00067287!). Elaphoglossum jinoteganum A.Rojas, Revista Biol. Trop. 51: 39. 2003. Type:—NICARAGUA. Jinotega: along Hwy 3, ca 1 km NW of La Fundadora entrance, unnamed peak ca 500 m W of Hwy, 13°01’N, 85°56’W, 1450–1520 m, 24 May 1981, W.D. Stevens & J. henrich 20411 (holotype: CR-106922!; isotype: MO-3326136!). Distribution:—Cuba, Jamaica, Haiti, Dominican Republic, Puerto Rico, Guadeloupe, Dominica, Mexico, Belize, Guatemala, Honduras, El Salvador, Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Ecuador, and Peru; 500– 3200 m. Remarks:—Fée (1845) cited three gatherings in the protologue of Acrostichum erinaceum, including one from Guadeloupe (de Thiouville s.n., P01818776), one from Mexico (Karwinski s.n., B-200070405), and one from Brazil (Gaudichaud s.n., B-200071871). This has caused confusion regarding the application of the name because each gathering represents a different species. Proctor (1977) designated the specimen from Guadaloupe as the lectotype, significantly narrowing the species concept. The other two specimens represent Elaphoglossum occidentale and E. scolopendrifolium, respectively, and should be disregarded as types. Typification aside, the E. erinaceum complex remains taxonomically difficult because of considerable variation in habitat, stem scale size and color, lamina shape and size, and degree of scaliness on the leaves. Much progress has been made in recent years regarding our understanding of this group (see Mickel & Smith 2004; Rojas-Alvarado 2003, 2010), but there is still a lot to be done. Some specimens from Mexico and Central America (e.g., Matos 2087 and Mickel 1083, NY) have extremely large leaves and tend to grow at higher elevations. As suggested by Mickel & Smith (2004), these may represent a distinct species to which the name E. fimbriatum T.Moore would apply. 5. Elaphoglossum glaziovii (Fée) Brade, Sellowia 18: 99. 1966. Acrostichum glaziovii Fée, Crypt. Vasc. Brés. 1: 6. 1869. Type:—BRAZIL. Rio de Janeiro: [Floresta da] Tijuca, [22°56’42’’S, 43°17’20’’W], 18 November 1867, A.F.M. Glaziou 2059 (lectotype, designated by Matos 2014: P-00249745!; isolectotypes: BR-0000006972530!, P-00249746!, P-00249747!, P-00249748!). (Figs. 2G, 3C, 10B) Distribution:—E Brazil, from Bahia to Rio Grande do Sul; 20–1800 m. Remarks:—This species is endemic to the Atlantic forest of eastern Brazil. It is characterized by having three kinds of scales on its sterile leaves: strongly subulate scales on petioles and costae, planar lanceolate scales along laminar margins, and branched microscales on both laminar surfaces. It is part of a species complex including the following species from eastern Brazil: E. amorimii, E. brevipetiolatum, E. prestonii, and E. scolopendrifolium (Matos & Mickel 2014). . Elaphoglossum gomezianum A.Rojas, Brenesia 45–46: 12. 1996. Type:—PANAMA. Chiriquí: Bugaba, Represa de Fortuna, Quebrada Honda, ca. 17.5 km N de los Planes de Hornito [8°28’48’’N, 82°37’12’’W], 1100 m, 3 May 1991, I.A. Valdespino & E. rios-Levy 1207 (holotype: NY-00039308!; isotypes: INB n.v., NY-4 sheets without number!, PMA-18142 [image!]). Distribution:—Costa Rica and Panama; 1000–1200 m. Remarks:—According to Rojas-Alvarado (1996a), the holotype of Elaphoglossum gomezianum is at NY, with duplicates at INB and PMA. There are five sheets of Valdespino & rios-Levy 1207 at NY, but only NY-00039308 has a pencil annotation that reads “holotype” in Rojas’ handwriting. This is also the only fertile specimen of this 30 • Phytotaxa 535 (1) © 2022 Magnolia Press MATOS ET AL. gathering and it was used for the protologue’s illustration. The other four sheets were found in the general collection with a printed label reading “Elaphoglossum gomezianum A.Rojas & Mickel – ined.”. They are now properly marked as isotypes of E. gomezianum A.Rojas. The INB isotype was not found during our visit and could be misfiled in the general collection. . Elaphoglossum hybridum (Bory) Brack., U.S. Expl. Exped. Filic. 16: 69. 1854. Acrostichum hybridum Bory, Voy. Îles Afrique 3: 95. 1804. olfersia hybrida (Bory) C. Presl, Tent. Pterid.: 235. 1836. Elaphoglossum hybridum (Bory) T. Moore, Index Fil. 10. 1857, nom. illeg. Type:—BOURBON [RÉUNION]. “Caverne Le Gentil, en allant au Piton des Neiges”, [21°05’54’’S, 55°28’51’’W], 2160 m, s.d., J. B. G. M. Bory s.n. (holotype: P-00466256!). (Figs. 10J–L, 11G) Acrostichum ciliare Carmich., Trans. Linn. Soc. London 12: 510. 1819, nom. illeg., non A. ciliare Thouars, Esquisse Fl. Tristan d’Acugna 32. 1804. Type:—TRISTAN DA CUNHA, [37°07’S, 12°17’W], [without day or month] 1817, D. Carmichael s.n. (lectotype, designated by Schelpe 1969: K-000369624!; isolectotype: BM-001066179!). Acrostichum ciliatum Desv., Ges. Naturf. Freunde Berlin Mag. Entdeck. Gesammten Naturk. 5: 310. 1811, nom. illeg., non A. ciliatum C. Presl, Reliq. Haenk. 1: 15. 1825. Type:—[RÉUNION]. “Habitat in insula Bourboniae”, s.d., P. Commerson s.n. (lectotype, designated by Schelpe 1969: P-JU 1000A n.v.; isolectotype: E-00429059!). Acrostichum hybridum Bory var. vulcani Lepervanche ex Fée, Mém. Foug. 2: 41. 1845. Elaphoglossum hybridum (Bory) Brack. var. vulcani (Lepervanche ex Fée) Christ, Neue Denkschr. Allg. Schweiz. Ges. Gesammten Naturwiss. 36: 104. 1899. Type:—BOURBON [RÉUNION]. Plaine des Palmistes, Plaine des Cafres, [21°10’S, 55°35’W], [without day or month] 1831–1836, M. LepervancheMézières s.n. (holotype: P-00466255!). Acrostichum melanopus Kunze, Linnaea 23: 214, 298. 1850. Elaphoglossum melanopus (Kunze) T.Moore, Index Fil.: 12. 1857. Type:— VENEZUELA. Caracas: [10°30’N, 66°55’W], s.d., G.K.W.h. Karsten s.n. (holotype: B-200071702!). Acrostichum propinquum Mett. ex Kuhn, Linnaea 36: 45. 1869. Elaphoglossum propinquum (Mett. ex Kuhn) Christ, Neue Denkschr. Allg. Schweiz. Ges. Gesammten Naturwiss. 36: 115. 1899. Type:—PERU. [Carabaya]: “prope Agapata” (Ayapata), [13°52’S, 70°19’W], June 1854, W. Lechler 2007 (lectotype, designated by Mickel 1991: B-200071702!; isolectotypes: K-sheet without number!, P00249940!). Acrostichum lindbergii Mett. ex Kuhn, Linnaea 36: 46. 1869. Elaphoglossum lindbergii (Mett. ex Kuhn) Rosenst., Hedwigia 46: 153. 1907. Type:—BRAZIL. [Minas Gerais]: Caldas, Chácara do Oliveira, [21°56’06”S, 46°25’24”W], January 1855, G.A. Lindberg 537 (lectotype, designated by Matos 2014: B-200071096!; isolectotypes: B-200071099!, K-sheet without number!). Acrostichum hybridum Bory var. minor Baker in Mart., Fl. Bras. 1(2): 574. 1870. Type:—BRAZIL. [Rio de Janeiro]: Serra dos Órgãos, [22°26’43”S, 42°59’59”W], March 1841, G. Gardner 5925 (Designated here: BM-sheet without number!; isolectotype: K-sheet without number!). Acrostichum tricholepis Baker, J. Bot. 29: 5. 1891. Elaphoglossum tricholepis (Baker) C.Chr., Index Filic.: 317. 1905. Type:— MADAGASCAR. Bekilus Mountains, [14°02’S, 48°19’E], December 1890, J.T. Last s.n. (holotype: K-000435725!). Elaphoglossum hybridum (Bory) Brack. var. cuneatum Bonap., Notes Pteridol. 14: 419. 1923. Type:—COMOROS. [Grande Comore]: “environs de Nioumbadjou”, [11°47’57”S, 43°17’50”E], 400–500 m, April 1911, M.L. humblot s.n. (holotype: P-00212873!). Elaphoglossum hybridum (Bory) Brack. var. cuneatum Bonap. f. minimum C.Chr., Notes Pteridol. 16: 128. 1925, nom. nud. Distribution:—South America (Argentina, Bolivia, Brazil, Colombia, Ecuador, Peru, Venezuela), islands of the midAtlantic Ocean (Gough Island, Inaccessible Island, Tristan da Cunha), Africa (Bioko, Cameroon, DRCongo, Ethiopia, Kenya, Malawi, Mozambique, Rwanda, South Africa, Tanzania, Uganda, Zimbabwe), Madagascar, and islands of the Indian Ocean (Comoros, Mauritius, Réunion); 50–4150 m. Remarks:—Elaphoglossum hybridum is the most widely distributed and probably the most variable species of E. sect. Polytrichia. The phenotypic plasticity of the sporophytes is overwhelming, even within small populations. Elaphoglossum hybridum var. vulcanii was described from a diminutive plant collected at high elevation on the exposed cliffs of a volcano. This phenotype is common throughout the range of E. hybridum, but there is always a complete series of intermediate forms between smaller and larger specimens from the same locality (see Matos & Mickel 2014, fig. 1). Christensen (1932) synonymized Elaphoglossum tricholepis under E. hybridum, but the yellowish-brown stem scales and the presence of subulate scales on laminar surfaces distinguish their types. Further studies may prove them to be distinct species. Roux (1982) noted that the type material of E. tricholepis was a mixed collection including a small individual of E. phanerophlebium C.Chr., but the sheet currently bears a handwritten label indicating that the admixture was removed. A NOMENCLATOR FOR ELAPhoGLoSSUM SECTION PoLYTrIChIA Phytotaxa 535 (1) © 2022 Magnolia Press • 3 The name Elaphoglossum hybridum (Bory) Brack. var. cuneatum Bonap. f. minimum was not validly published by Bonapart (1925) because it was not accompanied by a description or diagnosis (Turland et al. 2018, Art. 38.1). Several specimens from Madagascar (e.g., rakotondrainibe 2956, 5012, 5832, 6057, 6421, P; van der Werff 12863, 12895, MO) are unusual for having blackish resinous stems with lustrous reddish scales. These collections probably represent a distinct species, but additional studies are required. Specimens from Argentina, Bolivia, and southern Brazil tend to have yellowish to orangish stem scales and could also represent a distinct species. . Elaphoglossum hystrix (Kunze) T.Moore, Index Fil.: 10. 1857. Acrostichum hystrix Kunze, Linnaea 9: 26. 1834. Type:—PERU. “In sylvis densioribus fl. Peruv. ad Pampayaco”, [9°33’S, 75°52’W], July 1829, E.F. Poeppig s.n. (holotype: LZ, destroyed); Neotype:—PERU. “Peruv. subandin. in sylvis desioribus, Cuchero”, [9°30’S, 75°51’W], December 1829, E.F. Poeppig s.n. (neotype, designated here: W-0052695!). Distribution:—Peru and Bolivia; 1450–2200 m. Remarks:—The holotype of Acrostichum hystrix was at LZ (Mickel 1991), where the German botanist Kunze (1793–1851) used to work. Unfortunately, this herbarium was entirely destroyed in 1943 during World War II, leaving no extant original material (Thiers 2021). The neotype (W-0052695) was also collected by E.F. Poeppig in Peru, Huánuco, 1829, but the precise date and locality are different from those of the holotype (see above). . Elaphoglossum kessleri A.Rojas, Revista Biol. Trop. 50: 977. 2002. Type:—BOLIVIA. Cochabamba: José Carrasco Torrico, 137 km antigua carretera Cochabamba-Villa Tunari, 17°06’S, 65°34’W, 1700 m, 16 July 1996, M. Kessler, T. Krömer, J. Gonzales, B. hibbits & I. Jimenez 7345 (holotype: UC-1615246!; isotypes: LPB-0000065!, NY01402826!). Distribution:—Peru and Bolivia; 1700–2950 m. Remarks:—Previously, Elaphoglossum kessleri was known only from Bolivia (Rojas-Alvarado 2002), but it has now been collected in Peru (van der Werff 21399 p.p., MO, NY, UC). The subulate scales of this species are usually enrolled only at the base, not throughout their length as is typical of most subulate-scaled species. The scales thus appear lanceolate or ovate, not hair-like as in the other species. The name “E. nigroapiculatum Mickel & R.C.Moran, sp. nov. ined.”, which was annotated on several herbarium specimens but was never effectively published, refers to the fact that at least a few of the costal scales are blackened toward the apex. 30. Elaphoglossum latum (Mickel) Atehortúa ex Mickel, Fieldiana, Bot., n.s., 27: 123. 1991. Elaphoglossum apodum (Kaulf.) Schott ex J.Sm. var. latum Mickel, Amer. Fern J. 69: 100. 1979. Elaphoglossum latum (Mickel) Atehortúa ex Mickel, Novon 2: 377. 1992, nom. illeg., isonym. Type:—GUATEMALA. Alta Verapaz: along route 5 between Semococh and La Laguna, on road to Chajmayic, 15°30’N, 90°20’W, 500 m, 10 May 1942, J.A. Steyermark 46368 (holotype: F-1161805!; isotype: US-00048758!). Distribution:—Haiti, Dominican Republic, Mexico, Guatemala, Belize, Honduras, Nicaragua, Costa Rica, Panama, Colombia, and Ecuador; 0–1000 m (Matos et al. 2021). Remarks:—Elaphoglossum latum belongs to subsect. Apoda (Matos et al. 2021). Erik Leonard Ekman (1883– 1931) was probably the first to recognize the distinctness of this species from E. apodum, when he annotated some of his specimens from Hispaniola (e.g., Ekman 13406, B, K, NY, S) as E. apodum var. majus (nom. inval.). Christensen (1936) agreed that these specimens were exceptionally large for E. apodum, but considered that “size alone is insufficient as character of a species” and treated them under E. apodum. Many years later, Mickel (1979) described E. apodum var. latum Mickel. Atehortúa (1983) attempted to elevate this variety to species rank, but the name was not validly published because her doctoral thesis cannot be considered an effective publication (see Turland et al. 2018, Art. 30.9). Mickel (1991) validated E. latum Atehortúa ex Mickel in his treatment of the genus for Pteridophyta of Peru. Elaphoglossum latum does not occur in Peru, but the new combination was validly published under the comments of E. amazonicum (see Art. 41, Turland et al. 2018). Mickel (1992) accidentatlly published the same name again. 3. Elaphoglossum lonchophyllum (Fée) T.Moore, Index Fil.: 11. 1857. Acrostichum lonchophyllum Fée, Mém. Soc. Sci. Nat. Strasbourg 5(1): 68. 1857. Type:—MEXICO. Veracruz: Huatusco, [19°08’56”N, 96°57’58”W], May 1854, W. Schaffner 19 (holotype: RB-00671804!). (Figs. 3A, 3H) 3 • Phytotaxa 535 (1) © 2022 Magnolia Press MATOS ET AL. Elaphoglossum acutissimum Christ, Bull. Herb. Boissier, sér. 2, 5: 730. 1905. Type:—MEXICO. Chiapas: San Pablo, 1000–2000 m, [without day or month] 1904, G. Münch 168 (holotype: P-00603614!; isotypes: DS-267754 in CAS-0002516!, US-00067259!). Distribution:—Mexico, Honduras, Nicaragua, Costa Rica, and Panama; 600–1700 m (Matos et al. 2019). Remarks:—Elaphoglossum lonchophyllum belongs to the Decursivum group (Matos et al. 2019). The seven species of this group are unusual in sect. Polytrichia for lacking subulate scales and for having commisural veins. Elaphoglossum lonchophyllum closely resembles E. decursivum, but differs in having longer petioles (1/6–1/2 the leaf length vs. 1/10–1/5 the leaf length), darker petiolar scales, and laminae not long-decurrent. Also, the petiolar scales of E. lonchophyllum are composed of somewhat irregularly isodiametric or slightly elongated cells (Matos et al. 2019, fig. 5C). The holotype of Acrostichum lonchophyllum is at RB, as indicated by Windisch (1982). Three specimens (B200071831, B-200071833, and P-00603615) were also collected by Schaffner in Mexico, but their labels do not bear a collection number. The sheets at B were annotated by Mettenius as “Acrostichum schaffneri Mett.”, a name that was apparently never published, whereas the sheet at P was annotated by an unknown person as “Acrostichum lonchophyllum Fée”, with reference to the original publication. Since there is no evidence that these specimens were part of the original material, we do not regard them as types. In the protologue of Elaphoglossum acutissimum, Christ (1905) referred to a single collection (i.e., Münch 168) in his personal herbarium as the type (“typ. Herb. Christ”). Because his personal herbarium is now at P (Morton 1968), P00603614 is the holotype (not a syntype). Its original label was annotated in pencil by Christ as “n. sp.” 3. Elaphoglossum longipilosum (Atehortúa ex Mickel) A.Rojas, Brittonia 61: 299. 2009. Elaphoglossum auripilum Christ var. longipilosum Atehortúa ex Mickel, Novon 2: 369. 1992. Type:—COSTA RICA. Puntarenas: [Coto Brus], 5 km S of San Vito de Java, trail below Wilson Finca, [8°47’07’’N, 82°57’32’’W], 1130 m, 5 August 1967, r.M. Lloyd 4242 (holotype: NY-00149585!). Distribution:—Costa Rica and Panama; 500–2500 m (Matos et al. 2021). Remarks:—Elaphoglossum longipilosum belongs to subsect. Apoda (Matos et al. 2021). Its distinctness was first recognized by Atehortúa (1983), who proposed the name E. auripilum var. longipilosum Atehortúa (nom. inval.). This name, however, was not validly published because her doctoral thesis does not constitute effective publication (Turland et al. 2018, Art. 30.9). Mickel (1992) validated the varietal name, and Rojas-Alvarado (2009) elevated it to species rank (as recognized here and by Matos et al. 2021). 33. Elaphoglossum martinezianum A.Rojas, Amer. Fern J. 100: 174. 2010. Type:—MEXICO. Chiapas: SE side of Volcán Tacaná, above Talquian, [15°05’N, 92°05’W], 2200 m, 16 January 1973, D.E. Breedlove & A.r. Smith 31630 (holotype: MEXU-00881325!; isotypes: CAS-425112 [image!], NY-01402918!). Distribution:—Mexico, Guatemala, and El Salvador; 1550–2400 m. Remarks:—Until recently, Elaphoglossum martinezianum was known only from Mexico, but now also from Guatemala (Williams 26212, F) and El Salvador (Seiler 130, F, NY). A gathering from Ecuador (Luteyn 8408, NY, UC) may also represent this species, in which case E. martinezianum would be much more widespread than previously thought. 3. Elaphoglossum mexicanum (E.Fourn.) A.Rojas, Revista Biol. Trop. 51: 42. 2003. Acrostichum mexicanum E.Fourn., Mexic. Pl. 1: 63. 1872. Type:—MEXICO. [Puebla]: Barranca de Chinautla, [19°49’N, 97°22’W], 1981– 2133 m, s.d., F.M. Liebmann s.n (lectotype, designated here: P-00249876!; isolectotypes: GH-sheet without number!, K-sheet without number!). Other syntypes:—MEXICO. [Veracruz]: Misantla, [19°56’N, 96°51’W], June 1866, L. hahn s.n. (P-01405598!, P-01405599!). (Fig. 3E) Distribution:—Mexico, Guatemala, Honduras, and El Salvador; 1200–2700(–3300) m. Remarks:—Mickel & Smith (2004) placed Acrostichum mexicanum in the synonymy of Elaphoglossum erinaceum. The name was re-evaluated and transferred to Elaphoglossum by Rojas-Alvarado (2003), who compared it to E. barbatum, not E. erinaceum. The morphological differences between these species are few and subtle, but their distinctness has been supported by DNA evidence (Matos 2015). Elaphoglossum mexicanum has dark brown (vs. yellowish brown) stem scales, narrowly elliptic to oblanceolate (vs. broadly elliptic to oblong) sterile laminae, and more abundant scales on petioles, costae, and laminar margins. A NOMENCLATOR FOR ELAPhoGLoSSUM SECTION PoLYTrIChIA Phytotaxa 535 (1) © 2022 Magnolia Press • 33 35. Elaphoglossum mickeliorum F.B.Matos & R.C.Moran, Brittonia 69: 92. 2017. Type:—PERU. Cajamarca: San Ignacio, Dist. San José de Lourdes, localidad de Buenos Aires, Cerro El Parco, 05°42’04”S, 77°53’06”W, 1900 m, February 2002, r. Bonino 303 (holotype: MO-6105304!; isotypes: NY-02311231!, UC-1872427!). Distribution:—Endemic to Peru, in the states of Amazonas, Cajamarca, Cusco, and Pasco; 1700–2900 m (Matos & Moran 2017). Remarks:—Before being described as a new species, Elaphoglossum mickeliorum was commonly identified as E. erinaceum. It differs from all species of the E. erinaceum complex, however, by a nearly continuous band of flat (i.e., nonsubulate) scales along the margins of sterile laminae. In this character it is similar to E. amorimii, E. glaziovii, E. ornatum, E. prestonii, and E. scolopendrifolium, but it differs from these species by the presence of minute, erect, glandular hairs on the leaves (Matos & Mickel 2014, Matos & Vasco 2015). 3. Elaphoglossum miersii (Baker) C.Chr., Index Filic.: 311. 1905. Acrostichum miersii Baker, Ann. Bot. (Oxford) 5: 490. 1891. Type:—BRAZIL. Rio de Janeiro: [without precise locality], s.d., J. Miers s.n. (lectotype, designated by Matos et al. 2019: K-000727490!; isolectotype: BM-001027679!). (Fig. 2H) Elaphoglossum crispipalea M.Kessler & Mickel, Brittonia 58: 98. 2006. Type:—BOLIVIA. La Paz: Prov. Larecaja, Hacienda Simaco sobre el camino a Tipuani, 1400 m, February 1920, o. Buchtien 5176 (holotype: US-00955785!). Distribution:—Ecuador, Peru, Bolivia, and Brazil; 900–2800 m (Matos et al. 2019). Remarks:—Elaphoglossum miersii belongs to the Decursivum group, which is unusual in sect. Polytrichia for lacking subulate scales and for having commisural veins (Matos et al. 2019). Alston (1958) compared E. miersii to E. denudatum and E. lindbergii, being the first author to suggest that E. miersii could be closely related to other species of sect. Polytrichia. Elaphoglossum miersii most resembles E. papyraceum, which also occurs in Peru, Bolivia, and Brazil. In the field, both have spreading-arching sterile leaves and shorter, ascending to erect fertile ones. Elaphoglossum miersii differs by being smaller, with thinner stems (0.6–1.3 cm wide) and thinner petioles (1–2[–2.5] vs. 2.3–3.8 mm wide). The stem scales also differ: those of E. miersii are crispate with tortuous apices, whereas those of E. papyraceum are straighter and flatter. Additionally, the petioles of E. papyraceum bear dense microscales, which are sparse to absent in E. miersii. The isolectotype at BM, which was previously in Miers’s private herbarium, is fertile and bears the number “10.013,” which might refer to Miers’s collection number. 3. Elaphoglossum occidentale (Mickel) F.B.Matos, comb. et stat. nov. Elaphoglossum erinaceum var. occidentale Mickel, Mem. New York Bot. Gard. 88: 289. 2004. Type:—MEXICO. Mexico: about 6 miles south of Tenancingo and 8 miles north of Villa Guerrero on Route 55, [18°57’39”N, 99°35’26”W], 2600 m, 11 August 1960, J.T. Mickel & A.F. Clewell 706 (holotype: NY-00688052!; isotypes: ENCB n.v., MICH-1287229!, NY-sheet without number!, P01405634!, UC-1360344!). (Fig. 11H) Distribution:—W Mexico; 1400–2750 m. Remarks:—Elaphoglossum occidentale was originally described as a variety of E. erinaceum, from which it differs by orange-tan (vs. brown to black) scales that are often shorter (1–2 mm vs. 3–4 mm) on the sterile lamina margins. Also, E. occidentale is almost always terrestrial, whereas E. erinaceum is usually epiphytic. Several sheets of E. occidentale have been annotated as “E. morelosense A.Rojas”, but this name was never published. The same specimens from Mexico were later cited by Rojas-Alvarado (2003) as E. scolopendrifolium (Raddi) J. Sm. However, E. scolopendrifolium is endemic to eastern Brazil (Matos & Mickel 2014), whereas E. occidentale is restricted to western Mexico, from Sinaloa to Guerrero. The collections made by A. C. Sanders 4409 (UC) and 22572 (MO) in the Mexican state of Sinaloa are noteworthy for establishing the northern limit (ca. 23°N lat.) of section Polytrichia. 3. Elaphoglossum ornatum (Mett. ex Kuhn) Christ, Neue Denkschr. Allg. Schweiz. Ges. Gesammten Naturwiss. 36: 38. 1899. Acrostichum ornatum Mett. ex Kuhn, Linnaea 36: 44. 1869. Type:—VENEZUELA. Mérida: [8°36’N, 71°09’W], s.d., A. Engel 201 (holotype: B-200071399!). (Fig. 10C) Elaphoglossum heteroglossum Mickel, Ann. Missouri Bot. Gard. 78: 259. 1991. Type:—VENEZUELA. Portuguesa: Sucre, La Divisoria de la Concepción, 9°18’N, 70°06’W, 1500–1800 m, 23–26 October 1985, F. ortega, r. rivero, B. Stergios, h. van der Werff & G. Aymard 2748 (holotype: UC-1534033!; isotypes: MO-04903837!, NY-00149633!, NY-01402810!, PORT-16352!). 3 • Phytotaxa 535 (1) © 2022 Magnolia Press MATOS ET AL. Distribution:—NW Venezuela, restricted to the Andes; 400–1800 m. Remarks:—This name has been widely misapplied until recently. The species was originally described by Kuhn (1869) as Acrostichum ornatum Mett. ex Kuhn, based on a single specimen from Mérida, Venezuela. Upon transferring the species to Elaphoglossum, Christ (1899) considered A. amplissimum (=E. scolpendrifolium) as a synonym and expanded the geographic range of the species to Brazil. As a consequence, E. ornatum started to be widely used in floristic studies and herbarium labels of specimens from Brazil (mostly E. glaziovii) (Matos & Mickel 2014). In Venezuela, Vareschi (1969) misapplied the name E. ornatum to a species of sect. Lepidoglossa. This species, which was later described as E. fendleri F.B.Matos & A.Vasco, greatly differs from E. ornatum by the presence (vs. absence) of glandular hairs, ciliate (vs. entire) stem scales, planar and setose-margined (vs. subulate and entire) petiolar and costal scales, and setose-margined (vs. entire) lamina margin scales (Matos & Vasco 2015). Elaphoglossum ornatum is most similar to the Brazilian endemics E. amorimii, E. glaziovii, and E. prestonii, from which it differs by the more slender stems (up to 0.3 cm vs. 0.4–1.2 cm diam., without the scales) and lamina margin scales lanceolate with entire margins (vs. ovate to lanceolate with conspicuously toothed margins) (Matos & Mickel 2014). 3. Elaphoglossum papyraceum (Fée) F.B.Matos & R.C.Moran, Phytotaxa 353: 16. 2018. Acrostichum papyraceum Fée, Crypt. Vasc. Brés. 1: 4. 1869. Type:—BRAZIL. Rio de Janeiro: Serra do Lingua, 23 March 1869, A.F.M. Glaziou 2434 (lectotype, designated by Matos et al. 2019: P-00249910!; isolectotypes: BR-0000006970956!, P-00249909!). Acrostichum macrorhizum Baker, Ann. Bot. (Oxford) 5: 490. 1891. Elaphoglossum macrorhizum (Baker) C.Chr., Index Filic.: 310. 1905. Type:—BRAZIL. Rio de Janeiro: [precise locality unknown], s.d., J. Miers s.n. (holotype: K-sheet without number!). Elaphoglossum puberulentum M.Kessler & Mickel, Brittonia 58: 115. 2006. Type:—BOLIVIA. La Paz: Prov. Nor Yungas, Cantón Pacallo, Comunidad Chairo, Bajo Hornuni, subiendo 60 m altitudinales hacia el camino de la mina, 16°12’S, 67°53’W, 2000 m, 10 August 2002, S.G. Beck, K. Bach, A. Pareja & M. Villca 28386 (holotype: NY-00842485!; isotype: LPB-0000072 [image!]). Distribution:—Peru, Bolivia, and Brazil; 400–2520 m (Matos et al. 2019). Remarks:—Elaphoglossum papyraceum belongs to the Decursivum group (Matos et al. 2019). The seven species of this group are unusual in sect. Polytrichia for lacking subulate scales and for having commisural veins. Although typical subulate scales are lacking, most petiolar scales of E. papyraceum are patent and slightly enrolled at the base, revealing that they are actually subulate (Matos et al. 2019, figs. 4E). This also occurs in E. miersii, the most similar species morphologically, which also occurs in Peru, Bolivia, and southeastern Brazil (see discussion under that species). The cell shape in the petiole scales of the Decursivum group can be seen in fig. 5 of Matos et al. (2019). However, the caption of that figure is actually wrong: E. ambiguum should be 5A, E. decursivum 5B, E. lonchophyllum 5C, E. miersii 5D, E. papyraceum 5E, and E. vascoae 5F. Christ (1899) considered E. papyraceum a synonym of E. flaccidum (Fée) T. Moore [= E. rigidum (Aubl.) Urb.]. Alston (1958) recognized E. macrorhizum and E. miersii for Brazil, but also placed E. papyraceum as a synonym of E. rigidum. We now know that E. rigidum is a different species that belongs to sect. Elaphoglossum (Rouhan et al. 2004, Matos et al. 2019). Brade (1961) applied the name E. longifolium (Jacq.) J. Sm. to the Brazilian specimens of E. papyraceum, but the type of E. longifolium is an illustration (Plumier, 1705, t. 135) based on a currently lost specimen from Martinique. Matos & Mickel (2014) did not include E. papyraceum in their floristic treatment for Brazil because they were not aware that this species belonged to sect. Polytrichia. 0. Elaphoglossum pendulum A.Rojas, Brittonia 61: 295. 2009. Type:—COSTA RICA. Alajuela: Cantón de Alajuela, Distr. Sarapiquí, Colonia Virgen del Socorro, orillas del Río Sarapiquí, cerca de la nueva represa, [10°15’30’’N, 84°10’20’’W], 720 m, 27 December 2004, A. rojas & C. Frias 6309 (holotype: CR-0243085!; isotypes: K n.v., MO6378460!, NY n.v.). Distribution:— Costa Rica, known from less than 10 collections from Cordillera Central and Cordillera de Talamanca; 500–1400 m (Matos et al. 2021). Remarks:—Elaphoglossum pendulum belongs to subsect. Apoda (Matos et al. 2021). It has the longest leaves of all subulate-scaled species of Elaphoglossum, reaching up to 2 m long (Matos et al. 2021, fig. 1F). The long-pendent leaves with short petioles, linear laminae, and abundant, 0.3–0.7 mm long glandular hairs distinguish this species from other members of sect. Polytrichia. We did not find the isotypes at K and NY, even though they were cited in the protologue. A NOMENCLATOR FOR ELAPhoGLoSSUM SECTION PoLYTrIChIA Phytotaxa 535 (1) © 2022 Magnolia Press • 35 . Elaphoglossum polyblepharum Mickel, Ann. Missouri Bot. Gard. 78: 259. 1991. Type:—VENEZUELA. Amazonas: Atures, top of plateau, west side of valley of Río Coro-Coro, 8 km NNW of settlement of Yutaje, sandstone lajas and adjacent forest, W of Serranía de Yutaje, 5°41’00’’N, 66°08’30’’W, 900–1000 m, 1 March 1987, r. Liesner & B. holst 21517 (holotype: NY-00149695!; isotype: MO-1829741!, VEN-243906 [image!]). (Fig. 11J) Distribution:—Venezuela, Brazil, and Bolivia; 50–1060 m (Matos et al. 2021). Remarks:—Elaphoglossum polyblepharum belongs to subsect. Apoda (Matos et al. 2021). The species shows a curious disjunct distribution pattern, occurring in Venezuela (Mickel 1991), northwestern Brazil (Matos & Mickel 2014), and eastern Bolivia (Rojas-Alvarado 2002; Kessler et al. 2018). . Elaphoglossum prestonii (Baker) J.Sm., Ferns Brit. For. (ed. 2) 298. 1877, as “prestoni”. Acrostichum prestonii Baker, Gard. Chron. 1872: 1555. 1872, as “prestoni”. Acrostichum insigne Fée, Crypt. Vasc. Brésil 2: 6. 1873, nom. illeg. superfl. Type:—BRAZIL. Rio de Janeiro: “Itatiaia, prope Campo Bello”, [22°20’S, 44°35’W], 7 June 1871, A.F.M. Glaziou 5367 pro parte (lectotype, designated by Matos 2014: K-000994040!; isolectotypes: B-200071696!, BM-001073365!, K-000501504!, P-00249771!, P-00249772!, P-00249773!, P-00249774!, S-05-9352!). (Fig. 11I) Distribution:—SE Brazil; 1000–2100 m. Remarks:—Acrostichum prestonii was described partly from dried specimens collected by Glaziou in Brazil and partly from living material cultivated at Kew Gardens (Baker 1872). There are three sheets annotated as A. prestonii at K: two of them collected by Glaziou 5367 (K-000501504, K-000994040), and one undoubtedly vouchered from a cultivated plant (K-000501503). The cultivated plant, which was brought from Brazil by Rev. T.A. Preston, represents Elaphoglossum glaziovii and should be disregarded as a type. The other two specimens are syntypes of A. prestonii, from which Matos (2014) designated the lectotype. Working with a different set of Brazilian collections, Fée (1873) described Acrostichum insigne Fée based on the same type of A. prestonii Baker. By definition, A. insigne Fée is a superfluous name that must be rejected (Turland et al. 2018: Art. 52.1). Several sheets of Brade 8486 (GH, HB, NY, PH, R, SP, SPF) have been annotated as “Elaphoglossum insigne var. peroupavae” in Brade’s handwriting, but this varietal name was not published. See Matos (2014) and Matos & Mickel (2014) for additional comments. 3. Elaphoglossum procurrens (Mett. ex D.C.Eaton) T.Moore, Index Fil.: 365. 1862. Acrostichum procurrens Mett. ex D.C. Eaton, Filic. Wright. Fendler.: 194. 1960. Type:—CUBA. [Guantánamo]: “prope villam Monte Verde dictam, Cuba Orientali,” [20°19’N, 75°00’W], January–June 1859, C. Wright 793 (lectotype, designated by Matos et al. 2021: YU-000726!; isolectotypes: B-200071698!, BM-000605227!, E-00106045!, GOET-010306!, K-000590627p.p.!, Ksheet without number!, LE-00008867!, MA-695264!, MO-1867318!, NY-00127323!, P-01818731!, US-718540!, US-1411554!). Other syntypes:—CUBA. “In Cuba Orientali”, [without day or month] 1856–1857, C. Wright 793 (syntypes: BR-0000013093525!, K-000590627p.p.!, NY-00127325!, PH-611340!, YU-000724!); Cuba. “In Cuba Orientali”, [without day or month] 1859–1860, C. Wright 793 (syntypes: NY-00127324!, US-833950!, W-18890055843!, YU-000725!). (Fig. 2I) Distribution:—Cuba and Haiti; 150–1000 m. Remarks:—Eaton (1834–1895) described Acrostichum procurrens in his dissertation for Bachelor of Science degree, which was defended at Harvard University on July 14, 1860 (Eaton 1860). The manuscript was read to the American Academy of Arts and Sciences on August 14, 1860, and the dissertation was published on December 1860. A few months later, on February 21, 1861, this document was reprinted in the Memoirs of the American Academy of Sciences and Arts (Eaton 1861; exact date taken from Howard 1988). While consulting several online databases and printed indexes of plant names (e.g., Christensen 1906), we noticed a great amount of confusion regarding the dates, places, and even the authorities of the new names that were introduced by Eaton in his dissertation. Part of this confusion was probably caused because the text “Ex Mem. Acad. Amer. Scient. Et Artium, N. Ser. Vol. VIII.” was printed on the title page of Eaton’s dissertation. Also misleading is the entry for Eaton’s dissertation in Stafleu & Cowan (1976), which mistakenly states that Eaton (1861) was published in 1860. It is worth mentioning that dissertations published before 1 January 1953 constitute effective publication under the Shenzhen Code (Turland et al. 2018, Art. 30.9), and therefore A. procurrens was validly published in Eaton (1860). Also complicated was the lectotypification of this name. The process required a careful consideration of all problems associated with the Cuban collections of Charles Wright (see Howard 1988). We found 24 sheets of Elaphoglossum 3 • Phytotaxa 535 (1) © 2022 Magnolia Press MATOS ET AL. procurrens collected by Wright in no less than 13 herbaria. Three of these sheets (BM-000605226, K-000590628, and K-000590629) had no original labels. The others had a “Plantae Cubenses Wrightianae” label with number 793 on the upper left corner. According to Howard (1988) and references therein, this is a species number, not a collection number. Three kinds of labels were found: “Coll. C. WRIGHT, 1856–7, in Cuba Orientali” (label type 5 of Howard 1988: 86), “Coll. C. WRIGHT, prope villam Monte Verde dictam, Cuba Orientali, Jan.–Jul. 1859” (label type 2 of Howard, 1988: 86), “Coll. C. WRIGHT, in Cuba Orientali, 1859, 1860” (label type 4 of Howard, 1988: 86). There was no mention of a specific collection date in the protologue (Eaton 1860), so all of these specimens are syntypes. Adding to the complication, YU-000725 was annotated as holotype of A. procurrens by C. Sánchez in 2000. This specimen was formerly associated with YU-00024 and YU-000726, but the sheet was cut and each specimen was mounted on a separate sheet. Almost every other specimen had an isotype label. Matos et al. (2021) selected YU-000726 as the lectotype because it was fertile, complete (i.e., leaves attached to the stem), and represented by more duplicates that are widely distributed. Also, it has a small label printed with “Herb. D. C. Eaton”, suggesting that it was examined by Eaton (1860). . Elaphoglossum raywaense (Jenman) Alston, Bol. Soc. Brot., ser. 2, 32: 24. 1958. Acrostichum raywaense Jenman, Ferns Brit. W. Ind. 18: 341. 1909. Type:—GUYANA. Demerara: Mt. Raywa and upper parts of Isorooroo River, [6°48’N, 58°10’W], [without day or month] 1897, G.S. Jenman 1858 (lectotype, first step designated by Mickel, Fieldiana Bot., n.s. 27: 157. 1991, second step designated by Matos & Mickel, Brittonia 66: 389. 2014: NY-00214456!; isolectotype: NY-00214457!). Acrostichum apodum Kaulf. var. sprucei Baker in Mart., Fl. Bras. 1(2): 578. 1870. Type:—[BRAZIL]. [Amazonas: São Gabriel da Cachoeira], “prope San Gabriel da Cachoeira, ad Rio Negro, Brasiliae borealis”, [00°07’48’’S, 67°05’20’’W], January–August 1852, r. Spruce 2186 (lectotype, designated by Matos 2014: K-000898850!; isolectotypes: B-200071758!, B-200072410!, BM001026214!, BR-0000006972011!, CGE-13097 [image!], CGE-13098 [image!], CGE-13099 [image!], E-00911481 [image!], E00911486 [image!], GH-00521587!, K-000898849!, P-01464083!, P-01464084!, P-01464085!, US-00067318!, US-01100827!, W-1889-0200311 [image!]). Other syntypes:—[BRAZIL. Pará]: “Tanaii, ad Rio Acara, juxta Para”, [1°40’S, 48°25’W], July– September 1849, r. Spruce s.n. (syntypes: CGE-13096 [image!], K [on three sheets!], P-01464087!). Distribution:—Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Bolivia, and Brazil; 90– 1200 m (Matos et al. 2021). Remarks:—Elaphoglossum raywaense belongs to subsect. Apoda (Matos et al. 2021). It used to be confused with E. apodum from the Antilles, which differs by larger stem scales (5–28 × 0.3–1.5 vs. 5–8 × 0.2–0.5 mm) that are pinkish gold to ferrugineous, concolorous (vs. darker and resinous towards the apex, i.e., bicolorous), and conspicuously dentate (entire or nearly so). It is also similar to E. amazonicum, with which it is sympatric in Peru and Bolivia (see Matos et al. 2021 for a comparison). 5. Elaphoglossum reductum A.Rojas, Amer. Fern J. 100: 176. 2010. Type:—COSTA RICA. Cartago: Cantón de Turrialba, Reserva Indígena Chirripó, Cuenca del Matina, camino a Valle Escondido, entrado por las Brisas de Pacuarito, 9°56’40’’N, 83°28’40’’W, 750–850 m, 13 April 1999, A. rojas, L. Pacheco & E. Mora 5043 (lectotype, designated here: INB-0002841355; isolectotypes: MO-1079169 n.v., NY-01403008!). Distribution:—Costa Rica, known only from Fila Matama, on the Caribbean side of Cordillera de Talamanca; 720– 850 m. Remarks:—According to the protologue, the holotype of Elaphoglossum reductum should be at CR with duplicates at INB and MO. The specimen at INB has a note on its label stating that rojas et al. 5043 comprises 3 duplicates (dup.=3). The holotype was not found at CR. The isotype from MO was also not found, but we assume it is there because it has received barcode and accession numbers (http://www.tropicos.org/Specimen/2770162). To our surprise, we found a duplicate at NY. Because it is unlikely that a fourth duplicate exists, we here designate the specimen at INB as lectotype. No other gatherings are known for this species. . Elaphoglossum scolopendrifolium (Raddi) J. Sm., Bot. Mag. 72: 17. 1846. Acrostichum scolopendrifolium Raddi, Pl. Bras. Nov. Gen. 1: 4. 1825. Type:—BRAZIL. [Rio de Janeiro: Petrópolis, Fazenda Mandioca], [22°30’18’’S, 43°10’44’’W], s.d., G. raddi s.n. (lectotype, designated by Pichi Sermolli & Bizzarri 2005: PI-010927!; isolectotypes: BR-0000006972172!, FI n.v., PI-sheet without number [annotated by Pichi-Sermolli as 2]!). (Fig. 10E) A NOMENCLATOR FOR ELAPhoGLoSSUM SECTION PoLYTrIChIA Phytotaxa 535 (1) © 2022 Magnolia Press • 3 Acrostichum amplissimum Fée, Crypt. Vasc. Brés. 1: 5. 1869. Elaphoglossum amplissimum (Fée) Christ, Bull. Herb. Boissier, sér. 2, 2(3): 328. 1902. Type:—BRAZIL. Rio de Janeiro: [Angra dos Reis], Serra do Ariro, [22°53’52’’S, 44°20’33’’W], 19 June 1868, A.F.M. Glaziou 2436 (lectotype, designated by Matos 2014: P-00249820!; isolectotypes: P-00249821 p.p.!, P-00249822!, P-00249823!). Distribution:—SE Brazil; 40–1050 m. Remarks:—Acrostichum scolopendrifolium is one of the oldest names in Elaphoglossum sect. Polytrichia. In fact, only A. crinitum L. (1753) is older. Since it was described by Raddi (1825) from Brazil, the name has been widely applied for different subulate-scaled species throughout the neotropics (e.g., Sodiro (1883) for Ecuador; Shimek (1897) for Nicaragua; Rovirosa (1909) for Mexico; Domin (1929) for Dominica; Vareschi (1969) for Venezuela). The species, however, is endemic to southeastern Brazil (Matos & Mickel 2014). . Elaphoglossum semisubulatum R.C.Moran & Mickel, Brittonia 56: 203. 2004. Type:—BOLIVIA. La Paz: Larecaja, 40 km de Sorata a Quiabaya, [15°36’S, 68°36’W], 3500 m, 29 May 1995, M. Kessler, J. Gonzales, K. Bach & S. hohnwald 4311 (holotype: NY-00803976!; isotypes: AAU-sheet without number!, GOET n.v., LPB-0000074!, UC n.v.). (Fig. 10F) Distribution:—Bolivia; 2400–3500 m. Remarks:—Moran & Mickel (2004) cited isotypes of this species at AAU, GOET, LPB, and UC, but we did not find duplicates at GOET and UC. Elaphoglossum semisubulatum is known from less than 10 gatherings from Bolivia. It can be distinguished from other species in the section by the blade scales limited primarily to the costae and margins, bases of the sterile blades rounded to truncate, and presence of branched microscales on the abaxial surface of the laminae. Capitate-glandular hairs on the leaves are lacking. The blade scales are enrolled only at the base, not throughout their length as is typical of most subulate- scaled species. The scales therefore appear lanceolate, thus the specific epithet semisubulatum. This blade-scale characteristic is also found in E. kessleri, a Bolivian species that differs by narrowly cuneate laminae (vs. rounded to truncate) and scales of the costae and sometimes margins often darkened toward the apex (vs. concolorous). . Elaphoglossum silencioanum A.Rojas, Revista Biol. Trop. 51: 40. 2003. Type:—COSTA RICA. Puntarenas: Cantón de Coto Brus, Parque Internacional La Amistad, Cordillera de Talamanca, Valle del Silencio, sector de acampar a los jardines, 9°07’15’’N, 82°57’55’’W, 2500 m, 14 April 1996, F. Quesada, A. Mora, M. Moraga & A. rojas 1421 (holotype: INB-0001515453!; isotypes: CR n.v., MO n.v., NY n.v.). Distribution:—Costa Rica and Panama; 1100–2600 m. Remarks:—Elaphoglossum silencioanum belongs to the E. erinaceum complex. It differs from typical E. erinaceum by narrower stem scales (0.3–1 vs. 1–2.5 mm wide), longer petioles (2/5–1/2 vs. 1/3–2/5 of the leaf length), smaller laminae [14–31 × 3–5.5 vs. (14–) 20–40 (–65) × (4–) 5–9 (–11.5) cm], and absence of subulate scales on costae. The holotype has a printed note on its label that reads “dup.=3”, suggesting that this gathering was comprised of three duplicates. Rojas-Alvarado (2003) stated that these duplicates were at CR, MO, and NY, but we did not find any of them. . Elaphoglossum tambillense (Hook.) T.Moore, Index Fil.: 15. 1857. Acrostichum tambillense Hook., Icon. Pl. 7: pl. 656. 1844. Type:—ECUADOR. [Pichincha]: “on the sides of ravines at Tambillo, near Quito”, [0°25’S, 78°33’W], January 1843, W. Jameson s.n. (holotype: K-000590660!). (Fig. 3G) Acrostichum stramineum Mett., Ann. Sci. Nat., Bot., sér. 5, 2: 200. 1864, non Fée (1869). Elaphoglossum stramineum (Mett.) Christ, Neue Denkschr. Allg. Schweiz. Ges. Gesammten Naturwiss. 36: 117. 1899. Type:—COLOMBIA. Bogotá: Boqueron, [5°02’N, 74°33’W], 2700 m, s.d., A. Lindig 251 (lectotype, designated here: B-200072037!; isolectotypes: BM-000769787!, K-sheet without number!, NY-00149716!, NY-00149717!, P-00250021!, P-00250022!, P-00250023!, P-00250024!). Acrostichum pallidum Baker ex Jenman, J. Bot. 17: 263. 1879. Elaphoglossum pallidum (Baker ex Jenman) C.Chr., Index Filic. 312. 1905, non Hieronymus (1921). Type:—JAMAICA. [Without precise locality]: 1524 m, July 1878, G.S. Jenman 8 (lectotype, designated by Proctor 1985: K-000590610!; isolectotypes: E-00414654!, K-sheet without number!, NY-02008183!, NY-02008195 [fragment!], NY-02008199!). 3 • Phytotaxa 535 (1) © 2022 Magnolia Press MATOS ET AL. Distribution:—Jamaica, Haiti, Dominican Republic, Dominica, and Mexico to Ecuador; 1400–3900 m. Remarks:—There are two sheets annotated as isotypes of Acrostichum tambillense at P. Both specimens were collected by Jameson in Ecuador, but their labels do not agree with the holotype. They bear a collector’s number (n. 461) that was never mentioned in the protologue and their locality data was written in Latin, not English. Therefore, we do not consider these specimens as part of the original material. Elaphoglossum tambillense is extremely variable in leaf shape and size. It belongs to the E. hybridum complex and can be recognized by castaneous stem scales, minute glandular hairs on petioles and laminae, and subulate scales virtually lacking from the leaves. Juvenile plants may have conspicuous hydathodes on their sterile leaves, but field observations indicate that these structures eventually disappear with age. Some authors have recognized E. pallidum as a distinct species (e.g., Mickel 1995a; Mickel & Smith 2004), but we rather consider it a large phenotype of E. tambillense. Elaphoglossum stramineum also falls within the variation range of this species. Elaphoglossum tambillense var. major Mille (Rev. Col. Nac. Vicente Rocafuerte 10: 33. 1928) may be another synonym, but we could not find any original material to verify this. 50. Elaphoglossum trichophorum (Sodiro) C.Chr., Index Filic.: 317. 1905. Acrostichum trichophorum Sodiro, Anales Univ. Centr. Ecuador 19(135): 198. 1905. Type:—ECUADOR. Tungurahua: “Crescit in silv. subtrop. oriental. vulcani Tungurahua”, [1°28’01’’S, 78°26’30’’W], December 1904, L. Sodiro s.n. (lectotype, designated by Matos et al. 2021: QPLS-210822!; isolectotypes: B-200072182!, BM-001073359!, P-00522931!, P-sheet without number!, S-R-80!, SI006017 [image!], US-00067388!). (Fig. 3F) Distribution:—Endemic to the eastern slopes of the Andes in Ecuador; 1450–1800 m (Matos et al. 2021). Remarks:—Elaphoglossum trichophorum belongs to subsect. Apoda (Matos et al. 2021). The species is rare and endemic to Ecuador, being known from only eight gatherings, all sterile (i.e., without fertile leaves). Although B200072183 and P-00522930 have been recently annotated as types, they are not types because they were collected on a different date (i.e., October 1904 vs. December 1904). 5. Elaphoglossum ulei Christ, Neue Denkschr. Allg. Schweiz. Ges. Gesammten Naturwiss.: 114. 1899. Type:— BRAZIL. Santa Catarina: [Lages], “An einer felsenwand bei Minas”, [27°47’S, 49°29’W], June 1890, E.h.G. Ule 327 (lectotype, designated by Matos 2014: P-00249984!; isolectotypes: B-200070088 p.p.!, P-00249985!). (Fig. 11L) Elaphoglossum spannagelii Rosenst., Hedwigia 46: 153. 1907. Type:—BRAZIL. Santa Catarina: Lages, [27°47’S, 49°29’W], September 1904, P.C. Spannagel 2 (lectotype, designated by Matos 2014: S, accession S-R-1881!; isolectotypes: HB-40105!, HBR-39634!, R13771!, UC-443383!). Other syntypes:—BRAZIL. Santa Catarina: Lages, “ad ripas umbrosas flum. Rio do Caveiras”, [without day or month] 1905, P. C. Spannagel s.n. [Rosenstock, Filices austrobrasilienses, no. 3] (syntypes: B-200071967!, B-200072067!, BMsheet without number!, LE-00000073 [image!], LIL-001940 [image!], M-0206998!, NY-00149708!, NY-00149710!, P-00249968!, P00249969!, P-00249970!, P-00249971!, P-00249972!, P-00249973!, P-00249974!, R-109523!, S-05-9543!, US-532556!); BRAZIL. Paraná. Lucena [presently Santa Catarina, Itaiópolis], [without day or month] 1903, Wielewski s.n. (syntypes: NY-00149709!, S-117324!). Distribution:—S Brazil; 900–1030 m. Remarks:—Elaphoglossum ulei is a poorly known species from southern Brazil. It was known only from the state of Santa Catarina (Matos & Mickel 2014), but it has now been collected in Rio Grande do Sul (e.g., Gonzatti 2651, HUCS). Additional information on the nomenclature and taxonomy of this species can be found in Matos (2014) and Matos & Mickel (2014). 5. Elaphoglossum vascoae F.B.Matos & R.C.Moran, Ann. Missouri Bot. Gard. 48: 422. 2019. Type:—ECUADOR. Napo: Campamento de Inecel, cascada de San Rafael, [00°06’S, 78°35’W], 1400 m, 17 June 1996, h. Navarrete & P. Asimbaya 1767 (holotype: QCA-111528!; isotypes: AAU-sheet without number!, NY-02311333!). Elaphoglossum longifolium (Jacq.) J.Sm. var. pastazense Hieron., Hedwigia 48: 284. 1909. Type:—ECUADOR. Tungurahua: Pastaza valley, near Baños, s.d., M.A. Stübel 858 (holotype: B-200071198!). Distribution:—Colombia, Ecuador, and Peru; 750–2700 m (Matos et al. 2019). A NOMENCLATOR FOR ELAPhoGLoSSUM SECTION PoLYTrIChIA Phytotaxa 535 (1) © 2022 Magnolia Press • 3 Remarks:—Elaphoglossum vascoae belongs to the Decursivum group (Matos et al. 2019). Like the other species of this group, it is unusual in sect. Polytrichia for lacking subulate scales and for having commisural veins. Elaphoglossum vascoae most resembles E. ambiguum, E. lonchophyllum, and E. papyraceum, but differs by the petiole scales, which are weak and flaccid, tightly appressed, with obtuse apices, and composed of isodiametric cells. In contrast, the petiole scales of the other species are firm and spreading. Those of E. ambiguum are acuminate and composed of elongate (not isodiametric) cells; those of E. lonchophyllum are usually resinous and composed of irregularly shaped cells; and those of E. papyraceum are enrolled at the base. The cell shape in the petiole scales of the Decursivum group can be seen in fig. 5 of Matos et al. (2019). However, the caption of that figure is actually wrong: E. ambiguum should be 5A, E. decursivum 5B, E. lonchophyllum 5C, E. miersii 5D, E. papyraceum 5E, and E. vascoae 5F. 53. Elaphoglossum versatile (Sodiro) F.B.Matos, comb. nov. Acrostichum versatile Sodiro, Recens. Crypt. Vasc. Quit.: 76. 1883. Type:—ECUADOR. “En los bosques tropicales á la orilla del río Toachi”, s.d., L. Sodiro s.n. (holotype: currently lost). Neotype:—ECUADOR. [Pichincha]: “in silv. subtrop. valle de Mindo”, [0°03’04’’S, 78°46’41’’W], March 1900, L. Sodiro s.n. (neotype, designated here: P-sheet without number!; isoneotype: US-1196793!). (Figs. 5A–D, 11K) Distribution:—Colombia, Ecuador, and Peru; 1050–2700 m. Remarks:—We were not able to find the type specimen of Acrostichum versatile, even though we visited almost every herbaria known to have Sodiro’s original material (based on Stafleu & Cowan 1985). Among the seven gatherings annotated by Sodiro as A. versatile, there was not a single specimen from the type locality of Río Toachi, Ecuador. Moreover, the only collections made before 1883 (i.e., the year in which the name was published) were either made in Colombia (André 2374, 3606, NY) or do not present a precise locality (Sodiro s.n., UC-678152). Because it is impossible to determine if UC-678152 was part of the original material, we designate a neotype. Some morphological characters of E. versatile were discussed by Bell (1955), but he was not able to apply a name to the collection he used (Bell 909, GH, RB). Christ (1899) placed Acrostichum versatile in the synonymy of E. scolopendrifolium, which is endemic to eastern Brazil and differs by abundant subulate scales on petioles and costae (vs. virtually glabrous in E. versatile). Hybrids Elaphoglossum × morphohybridum A.Rojas [Elaphoglossum alvaradoanum A.Rojas × E. crinitum (L.) Christ], Open Access Libr. J. 6 (e5172): 4. 2019, new name for Elaphoglossum × intermedium A.Rojas, Acta Bot. Malac. 42: 96. 2017, nom. illeg., non E. intermedium Brack., U.S. Expl. Exped., Filic. 16: 69. 1854. Type:—COSTA RICA. Puntarenas: Isla del Coco, Parque Nacional Isla del Coco, Meseta Occidental, en los nacientes de Quebrada Albatros, [05°31’15”N, 87°04’40”W], 350–400 m, 1 January 2011, A. rojas & C. herrera 9281 (holotype: CR; isotype: MO-4933293, currently lost). Distribution:—Cocos Island, a volcanic island about 500 km SW of mainland Costa Rica; 350–400 m. Remarks:—Elaphoglossum × morphohybridum is a putative hybrid between E. alvaradoanum and E. crinitum, and its morphology is intermediate between these species. Both parental species belong to E. sect. Polytrichia, being characterized by the presence of subulate scales and lack of hydathodes, but only E. alvaradoanum belongs to subsect. Apoda. Elaphoglossum crinitum is probably the most distinctive species of the genus, being characterized by long petioles (ca. 1/2 of sterile leaf length), broadly elliptic laminae, rounded laminar bases and apices, and regularly anastomosing veins. The other parent, E. alvaradoanum, has short petioles [(1/5–) 1/10–1/20 of sterile leaf length], oblanceolate laminae, attenuate to cuneate laminar bases, acuminate to caudate laminar apices, and free veins. The hybrid has petioles of intermediate length (ca. 1/4 of sterile leaf length), broadly oblanceolate laminae, cuneate laminar bases, obtuse laminar apices, and partially anastomosing veins. Unfortunately, E. × morphohybridum is known from a single gathering comprising two sterile specimens. Elaphoglossum × setaceum Lorence [Elaphoglossum hybridum (Bory) Brack. × E. lanatum (Bojer ex Baker) Lorence], Fern Gaz. 12(6): 348. 1984. Type:—MAURITIUS. Tamarind Falls Reservoir, on west bank of small river just north of the reservoir, [20°21’20’’S, 57°27’22’’E], 4 April 1973, D.h. Lorence 15823b (holotype: MAU-0013385!). (Fig. 12) Distribution:—Mauritius; ca. 500 m. Remarks:—Elaphoglossum × setaceum was described based on a single sterile individual from Mauritius. According to the protologue, the hybrid was found amongst a colony of E. lanatum (Bojer ex Baker) Lorence 0 • Phytotaxa 535 (1) © 2022 Magnolia Press MATOS ET AL. (Lorence 15823c, MAU) that was growing on a shady stream bank, directly below a vigorous colony of E. hybridum (Lorence 15823a, MAU). No other members of the genus occurred in the vicinity, and the hybrid was morphologically intermediate between those two species. The holotype sheet contains three specimens (i.e., the hybrid and its putative parents), each with a different barcode number. The original label has a drawing that shows the ecological condition in which they were found (Fig. 12). Elaphoglossum lanatum belongs to sect. Lepidoglossa (Rouhan et al. 2004), meaning that this is the only reported case of hybridization between sections in Elaphoglossum. Doubtful names and excluded species Elaphoglossum cinctum Rosenst., Repert. Spec. Nov. Regni Veg. 25: 63. 1928. Type:—BOLIVIA. [La Paz: Larecaja], San Carlos, near Mapiri, 800 m, 12 February 1927, o. Buchtien 301 (lectotype, designated here: S, no. R-1842; isolectotype: UC-478103!). Rojas-Alvarado (2002) suggested that this species was part of Elaphoglossum sect. Polytrichia, but the type specimens of E. cinctum represent juvenile plants of olfersia cervina (L.) Kunze (Dryopteridaceae) (Kessler et al. 2018). Indeed, these plants are similar to some species of the Decursivum group (sensu Matos et al. 2019), especially because of their simple entire leaves with commissural veins. However, the long-creeping stems with two rows of leaves, the lack of an elongated ventral meristele as seen in a cross-section of the stem, and the absence of phylopodia are characteristics of olfersia. Elaphoglossum demissum Christ, Bull. Soc. Bot. Genève, sér. 2, 1(5): 222. 1909. Type:—COSTA RICA. [San José]: La Palma, 1500 m, [without day or month] 1930, C. Wercklé 180 (lectotype, designated here: P-00602734 p.p.!). Elaphoglossum demissum was considered a synonym of E. erinaceum by Mickel (1995a). It was described from a mixed herbarium sheet (P-00602734) containing four specimens representing two species. The original description was almost entirely based on the single fertile specimen of that sheet. Having long creeping stems, virtually glabrous leaves, and ovate sterile laminae, this fertile specimen clearly belongs to E. sect. Elaphoglossum and is here designated as the lectotype of E. demissum. The other three sterile specimens are juvenile plants of E. angustioblongum. Christ (1909) used them only for the description of the veins, which supposedly end in hydathodes (“nervis […] in hydathodas nigras terminantibus”). Elaphoglossum lalitae L.D.Goméz in L.D.Gómez & J.G.Laurito, Phytologia 60: 74. 1986. Type:—COSTA RICA. Limón: Asunción, Río Segundo, 400–500 m, 28 April 1985, J. Berrocal, A. Solís, G. herrera & J. rodríguez 43 (holotype: CR-113182). Gómez & Laurito (1986) placed this species in sect. Decorata Mickel & Atehortúa, suggesting a close relationship to Elaphoglossum decoratum. These two species are superficially similar in laminar shape and petiolar indument (the petioles are densely covered by broad, light-colored scales), but they are not closely related. A recent molecular phylogenetic study (Matos et al. 2019) confirmed that E. decoratum is nested within section Polytrichia, whereas E. lalitae belongs to E. sect. Elaphoglossum. A re-examination of specimens provided additional differences between the two species: E. lalitae has narrowly winged spores (vs. broadly folded), petiolar scales with isodiametric cells (vs. at least twice as long as wide, with strongly sinuate anticlinal walls to form a jigsaw pattern), and sterile laminae without marginal scales (vs. marginal scales present, forming a nearly continuous row around the lamina). Elaphoglossum lucens A.Rojas, Amer. Fern J. 100: 172. 2010. Type:—PERU. Cajamarca: San Ignacio Province, Distrito Huarango, El Convento, 1100–1400 m, 2 July 1996, J.C. de la Cruz & E. rodríguez 2866 (holotype: MO-4933293, currently lost). Elaphoglossum lucens was described from a single specimen collected in Peru. The holotype has received both accession and barcode numbers at MO (http://www.tropicos.org/Specimen/1204896), but it was not found in that herbarium. We refrain from using this name until the holotype is found. Elaphoglossum neeanum A.Rojas, Revista Biol. Trop. 51: 40. 2003. Type:—PANAMA. Darién: middle slopes on W side of Cerro Pirre, 800–1050 m, 29 June 1988, T.B. Croat 68931 (holotype: MO-3618868, currently lost). Elaphoglossum neeanum was described with other species of E. sect. Polytrichia from Mexico and Central America (Rojas-Alvarado 2003). It was compared to E. cinctum (=olfersia cervina, see above) and E. albomarginatum, which A NOMENCLATOR FOR ELAPhoGLoSSUM SECTION PoLYTrIChIA Phytotaxa 535 (1) © 2022 Magnolia Press • is one of the few species of sect. Polytrichia without subulate scales. We did not find the holotype of E. neeanum at MO, but none of the paratypes available for examination (Atehortua 1169, MO; Pipoly et al. 16855, MO) had subulate scales. In contrast to other glabrous species of sect. Polytrichia (such as E. albomarginatum and others of the Decursivum group), E. neeanum probably belongs to E. sect. Elaphoglossum because it has long-creeping stems, distichous leaves, and no glandular hairs. A careful examination of the holotype would help to confirm the exclusion of this species from sect. Polytrichia. Elaphoglossum siliquoides (Jenman) C.Chr., Index Filic.: 315. 1905. Acrostichum siliquoides Jenman, J. Bot. 19: 53. 1881. Type:— JAMAICA. [Without precise locality]: 610–915 m, March 1877, G.S. Jenman 6 (holotype: K-001089661!). This species was placed in subsect. Apoda by Mickel & Atehortúa (1980), but the presence of hydathodes on the leaves of adult sporophytes indicates that it does not belong in this group. Molecular phylogenetic studies based on plastid DNA data have recently confirmed that E. siliquoides belongs to the Setosa Clade, being more closely related to E. costaricense Christ (Rouhan et al. 2004; Matos et al. 2019). Elaphoglossum solomonii A.Rojas, Revista Biol. Trop. 50: 983. 2002. Type:—BOLIVIA. La Paz: Murillo Province, Valle del Río Zongo, 36.7 km al N (abajo) de la cumbre, ca. 2 km al N de La Planta Hidroeléctrica Chururaqui, 1800 m, 30 April 1990, J.C. Solomon 18827 (holotype: UC-1584521!; isotypes: LPB-0000075!, MO-3896498!). Elaphoglossum solomonii is endemic to Bolivia (Kessler et al. 2018). It was placed in section Polytrichia by Rojas-Alvarado (2002), who overlooked the presence of conspicuous hydathodes and compared it to E. cordifolium (=E. barbatum) in the Latin diagnosis and discussion. Based on the presence of subulate scales and hydathodes, it belongs to sect. Setosa. Elaphoglossum squamicostatum A.Rojas, Brenesia 47–48: 13. 1997 [1998], as “squamocostatum”. Type:—COSTA RICA. Cartago: Paraíso, Valle del Reventazón, Orosi, Estación de Biología Tropical Río Macho y alrededores, 9°46’N, 83°52’W, 1550 m, 7 April 1994, A. rojas 1104 (holotype: INB-1593336 [image!]; isotypes: CR-198318 n.v., MO-5886493 n.v.). Elaphoglossum squamicostatum is a poorly known species from the West Indies, Guatemala, Honduras, Costa Rica, Panama, and Ecuador (Rojas-Alvarado 2002). Its phylogenetic position is unknown, as it has never been included in a molecular phylogenetic study. Morphologically, it is similar to E. vascoae by nearly glabrous leaves, appressed petiolar scales composed of isodiametric cells, thin-textured laminae, and presence of a commissural vein. Based on these characters, we suspect that E. squamicostatum might belong to the Decursivum group of E. sect. Polytrichia (sensu Matos et al. 2019). However, it differs from the species of that group by the presence of large and persistent ovate scales along the costae of sterile leaves. This is either an apomorphy within the Decursivum group or an indication that E. squamicostatum actually belongs to sect. Elaphoglossum. Elaphoglossum webbii (Bory ex Fée) T.Moore, Index Fil.: 16. 1857. Acrostichum webbii Bory ex Fée, Mém. Foug., 2. Hist. Acrostich.: 51, tab. 24, fig. 4 [erroneously cited as “pl. XXII”]. 1845. Type:—PANAMA. [Without precise locality], [without day or month] 1833, Webb s.n. (lectotype, designated by Rouhan & Cremers 2006: P-00250049!). Acrostichum webbii was considered a synonym of Elaphoglossum apodum by Christ (1899), Christensen (1906), and Rouhan & Cremers (2006). The lectotype of A. webbii, however, represents E. setigerum (Sodiro) Diels, a species that belongs to E. sect. Setosa (Christ) Mickel & Atehortúa and differs from E. apodum by longer petioles (1/4–1/2 vs. less than 1/30 of sterile leaf length) and presence of hydathodes. Moreover, E. apodum is endemic to the Antilles, whereas the lectotype of A. webbi is from Panama. To promote nomenclatural stability, Matos (2020b) proposed the rejection of A. webbi under Art. 56 of the Shenzhen Code (Turland et al. 2018). Acknowledgements This paper is part of the doctoral dissertation of FBM, which was done under the supervision of Robbin C. Moran, in the joint Graduate Studies Program of the New York Botanical Garden and City University of New York. We thank the dissertation committee members Alejandra Vasco, Amy Berkov, Fabian Michelangeli, John Mickel, and Joseph • Phytotaxa 535 (1) © 2022 Magnolia Press MATOS ET AL. Rachlin for comments and corrections on the first draft of this manuscript. Major funding for this study was provided by CAPES, CNPq, Fulbright, The City University of New York, and The New York Botanical Garden. Support for fieldwork in Brazil, Costa Rica, Ecuador, and Mexico was provided by CAPES/Fulbright (PhD scholarship to FBM), The Organization for Tropical Studies (OTS Teaching Assistantship to FBM), and The International Association for Plant Taxonomy (IAPT Research Grant to FBM). FBM also acknowledges the American Fern Society (AFS Travel Award 2011, 2013), the American Society of Plant Taxonomists (ASPT Travel Award 2011, 2013), and The Smithsonian Institution (Cuatrecasas Award 2010) for helping with funds for herbarium studies. We thank the curatorial staff of the following herbaria for making their specimens available: AAU, B, BHCB, BM, BOLV, BR, BSC, CAS, CEPEC, COL, CR, F, FLOR, FURB, GH, HAC, HAJB, HB, HBG, HBR, HLDG, HUA, HUEFS, HULE, INB, INPA, JAUM, JBSD, K, LIL, LPB, LSCR, M, MA, MBM, MEXU, MGC, MICH, MO, MSC, NY, P, PH, PI, Q, QCA, QCNE, QPLS, R, RB, S, SEL, SP, SPF, UC, ULV, UPCB, US, and YU. Alan Smith (UC) was particularly helpful to FBM during a visit to California. 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(1966) A cytotaxonomic survey of the pteridophytes of Jamaica. Earth and Environmental Science Transactions of the royal Society of Edinburgh 66 (9): 169–237. https://doi.org/10.1017/S0080456800023516 Walker, T.G. (1985) Cytotaxonomic studies of the ferns of Trinidad 2. The cytology and taxonomic implications. Bulletin of the British Museum (Natural history), Botany series 13 (2): 149–249. Windisch, P.G. (1982) Specimens from Fée’s pteridological collection at the Botanical Garden of Rio de Janeiro. American Fern Journal 72: 56–60. https://doi.org/10.2307/1547061 Zona, S. & Christenhusz, M.J.M. (2015) Litter-trapping plants: filter-feeders of the plant kingdom. Botanical Journal of the Linnean Society 179: 554–586. https://doi.org/10.1111/boj.12346 APPENDIX . Numerical List of Taxa and Specimens Examined. Numerical List of Taxa 1. Elaphoglossum albomarginatum 2. Elaphoglossum alvaradoanum 3. Elaphoglossum amazonicum 4. Elaphoglossum ambiguum 5. Elaphoglossum amorimii 6. Elaphoglossum angustioblongum 7. Elaphoglossum apodum 8. Elaphoglossum atehortuae 9. Elaphoglossum auripilum 10. Elaphoglossum backhouseanum 11. Elaphoglossum baquianorum 12. Elaphoglossum barbatum 13. Elaphoglossum beitelii 14. Elaphoglossum blepharoglottis 15. Elaphoglossum brevipetiolatum 16. Elaphoglossum clathratum 17. Elaphoglossum cotoi 18. Elaphoglossum crinitum 19. Elaphoglossum crispatum 20. Elaphoglossum cubense 21. Elaphoglossum decoratum 22. Elaphoglossum decursivum 23. Elaphoglossum denudatum 24. Elaphoglossum erinaceum 25. Elaphoglossum glaziovii 26. Elaphoglossum gomezianum 27. Elaphoglossum hybridum 28. Elaphoglossum hystrix 29. Elaphoglossum kessleri 30. Elaphoglossum latum 31. Elaphoglossum lonchophyllum • Phytotaxa 535 (1) © 2022 Magnolia Press MATOS ET AL. 32. Elaphoglossum longipilosum 33. Elaphoglossum martinezianum 34. Elaphoglossum mexicanum 35. Elaphoglossum mickeliorum 36. Elaphoglossum miersii 37. Elaphoglossum occidentale 38. Elaphoglossum ornatum 39. Elaphoglossum papyraceum 40. Elaphoglossum pendulum 41. Elaphoglossum polyblepharum 42. Elaphoglossum prestonii 43. Elaphoglossum procurrens 44. Elaphoglossum raywaense 45. Elaphoglossum reductum 46. Elaphoglossum scolopendrifolium 47. Elaphoglossum semisubulatum 48. Elaphoglossum silencioanum 49. Elaphoglossum tambillense 50. Elaphoglossum trichophorum 51. Elaphoglossum ulei 52. Elaphoglossum vascoae 53. Elaphoglossum versatile Specimens Examined Specimens are arranged by collector (with first initial, when known) in alphabetical order, followed by collector’s number in increasing order (s.n. = without number), followed by species number in parentheses. Boldface indicates types. Abbott, W. 276 (7) Abbott, W. 2072 (18) Acebey, A. 425 (44) Acevedo-Rodríguez, P. 9008 (44); 12825, 13020, 13360 (49); 13782 (21); 13910, 14235 (18) Acuña, J. 9965, s.n. [HAC-SV23790], s.n. [HAC-SV9965], s.n. [ULV-SV9965] (20) Adamson, G. 389 (27) Aguilar, R. 2741 (32); 5076 (17) Aguinda, R. 849 (12) Alain , Bro. 3264, 5170, 5453 (7); 247 (20) Alem, M. 187, 202 (41) Alfaro, E. 612 (24); 1464 (32) Allard, H. 17400 (49); 20888 (3) Allen, P. 233 (30), 1896 (18) Alluaud, M. s.n. [P-00605516] (27) Almeida-Neto, M. 127 (46) Almeida, T. 2570, 2621 (44); 2837 (27); 2855 (25); 5319 (24); 5525, 6435 (4); 6854 (24); 7196, 7440 (12); 7899 (52); 8368 (13); 8990, 9007 (25) Altamirano, S. 734, 1208, 1073 p.p. (41); 1256-b (36) Álvarez , A. s.n. [HAJB-HFC57453] (7); s.n. [HAJB-HFC64587] (20) Amorim, A. 4248 (25) Anderson, W. 3139 (7); 3141 (18); 3142 (23); 3154 (49); 3304 (23); 3307 (7); 3317 (23); 3318 (7); 3442 (49); 4449 (37); 4728 (12); 4729 (24); 7522 (27) Andrade, I. 838 (25) André, E. 2374, 3606 (53) Andriamihajarivo, T. 124 (27) Angulo, L. 605 (18) A NOMENCLATOR FOR ELAPhoGLoSSUM SECTION PoLYTrIChIA Phytotaxa 535 (1) © 2022 Magnolia Press • Ankersen, H. 103 (52); 153 (53) Anonymous s.n. [K-000500] (10) Antonio, T. 1094 (1) Araújo, I. 255 (44) Arbeláez, A. 315 (21) Areces , A. s.n. [HAJB-HFC25739] (7) Arias, I. s.n. [HAJB-HFC50025] (7) Ariati, V. 42, 187, 383, 1081 (25) Ariste-Joseph, Bro. A353 (12) Arroyo, L. 810 (41) Arruda, A. 4 (25); 47, 112 (27); 130 (25) Arsène, G. 6097, 8473, s.n. [P-01464590], s.n. [P-01624413], s.n. [Rosenstock, Filices Mexicanae no. 62] (37) Aschenbach, F. s.n. [P-01380564] (46) Ash, J. 3383 (27) Assi, L. 16341 (18) Athayde, F. 119, 229, 266 (25) Aulestia, M. 45, 160, 303, 2202 (44) Aymard, G. 967 (44) Azofeifa, A. 76 (24); 423 (30) Bach, K. 1490 (27); 1571, 1573 (39); 1866 (12) Badini, J. 29 (27); 34, s.n. [OUPR-10150], s.n. [OUPR-10153], s.n. [OUPR-10154], s.n. [OUPR-10155], s.n. [OUPR-10157], s.n. [OUPR10159], s.n. [OUPR-10161], s.n. [OUPR-10162] (21); s.n. [OUPR-10758], s.n. [OUPR-10759], s.n. [OUPR-10761], s.n. [OUPR-10762], s.n. [OUPR-10763] (27) Baker, M. 6887 (44); 7316 (12) Balch, A. s.n. [NY-02008008] (7) Balslev, H. 4672 (30); 84492 (44) Bang, M. 2608 (12) Baracaldo, P. 69 (12) Barbosa, E. 3453 (25) Barbour, P. 2757, 2847 (12); 4175 (27); 2528-a (35); 2528-b (27) Barcia, J. 905 (25) Barnett, L. 330-b (27) Barros, F. 790, 929 (25) Barros, W. 693 (42) Bartholomew, B. 2968 (37) Bass, M. 97, 149, 169, 333 (30) Bässler, M. s.n. [HAJB-HFC60627], s.n. [HAJB-HFC60940], s.n. [HAJB-HFC60963-A] (7) Batista, J. 912 (10); 1258 (21); 1496 (18) Baudouin, A. 804 (27) Beck, S. 504 (12); 586 (27); 3117, 3150 (29); 4056, 4929 (12); 7382 (36); 7599, 11964, 11967 (27); 18478, 22858 (12); 28384 (29); 3 (39); 28458 (29); 28520 (36) Bécquer, E. s.n. [HAJB-HFC82891] (7) Beitel, J. 533 (13) Bélanger, C. s.n. [P-01409004] (18) Bell, P. 272 (30); 434, 441, 528 (52); 568 (12); 855 (50); 909 (53) Belo, W. 517 (25) Beltran, H. 525 (44) Benl, G. FP124 (27) Bequaert, Dr. 4642, s.n. [P-00605518] (27) Bernal, H. 62 (30) Bewsher, C. s.n. [US-03339952] (27) Bisse, J. 5114 (7); 39648 (18); 50025, s.n. [HAJB-HFC14750], s.n. [HAJB-HFC25720-A], s.n. [HAJB-HFC25795], s.n. [HAJB-HFC2708], s.n. [HAJB-HFC27608], s.n. [HAJB-HFC36937], s.n. [HAJB-HFC37003], s.n. [HAJB-HFC37182-A], s.n. [HAJB-HFC37634] (7); s.n. [HAJB-HFC37707] (20); s.n. [HAJB-HFC39723], s.n. [HAJB-HFC39808], s.n. [HAJB-HFC41050], s.n. [HAJB-HFC45304-A], s.n. 50 • Phytotaxa 535 (1) © 2022 Magnolia Press MATOS ET AL. [HAJB-HFC45308], s.n. [HAJB-HFC45373], s.n. [HAJB-HFC46238], s.n. [HAJB-HFC49466], s.n. [HAJB-HFC49739], s.n. [HAJBHFC5114], s.n. [HAJB-HFC5162], s.n. [HAJB-HFC52125], s.n. [HAJB-HFC52281], s.n. [HAJB-HFC52286], s.n. [HAJB-HFC52305], s.n. [HAJB-HFC53215], s.n. [HAJB-HFC53285], s.n. [HAJB-HFC53389] (7) Bittner, J. 161 (12); 420 (18); 1041 (24); 1045 (18); 1452 (22); 1862 (24) Bittner, J. 2152 (18); 2499 (12) Blackmore, S. 386 (27) Bodenghien, A. 1313, 1848 (27) Bohs, L. 1757 (34) Boiteau, M. 3003 (27) Boivin, L. s.n. [P-00605461] (27) Bonino, E. 744 (28); 1145 (39); 1183 (28) Bonino, R. 184 (35); 290 (52); 293-a (35); 297 (53); 303 (35); 327 (27); 365, 451 (44) Borges, R. 527 (25) Borgo, M. 2472 (25) Bory, J. s.n. [Fougères catal. No. 37-23], s.n. [Herbar. Filic. Luerssen, n. 9] (18); s.n. [P-005], s.n. [P-00605455] (27) Bourgeau, M. 2888 (31) Boutin, F. 2570 (37); 5071 (30) Box, H. 274 (18); 278 (7); 492 (18); 501 (7) Boyle, B. 4505, 5012 (12); 5820 (6); 6867 (17) Brade, A. 132 (24); 185 (31); 276 (9); 523 (24); 5024 (25); 5288 (21); 6555 (42); 7606 (27); 8388, 8390 p.p. (25); 8390 p.p. (46); 8391 (39); 8480 (25); 8486 (42); 8595, 8613 (25); 8632 (46); 9292, 9381 (27); 9426 (25); 9427 (27); 9548, 9555 (46); 9656 (27); 9728, 9819 (25); 9870 (27); 10246 (42); 10253 (25); 10388 (42); 10808 (27); 10868 (25); 11536 (27); 11645 (25); 12080 (46); 14056 (21); 14933, 14934 (46); 15207 (25); 15514 (42); 15515, 15516, 16284 (27); 16404 (39); 16465, 16725 (25); 16813 (9); 17101 (27); 17165 (39); 17348 (46); 17973 (27); 19244 (42); 19421 (21); 21117, s.n. [HB-40076], s.n. [HB-40077] (25); s.n. [HB-54230] (27); s.n. [NY-01015658], s.n. [NY-01015669] (25); s.n. [R-86270] (46); s.n. [R-86271], s.n. [RB-01075038] (25); s.n. [RB-01075041] (21) Braithwaite, A. 420, 453 (27) Brandbyge, J. 32531, 32577 (44) Brass, L. 16302, 16662 (27) Breedlove, D. 15222 (34); 21510, 22347, 22410 (24); 22804 (49); 23194, 27709 (24); 27709 (34); 29815 (24); 3 (1); 330 (33); 31804 (24); 32046 (49); 32209 (24); 32218 (22); 40339 (49); 48328 (30); 57355 (30) Brenes, A. 4134 (6); 4181 (21); 4917 (24); 14262 (6); 15099 (31) Brina, A. s.n. [BHCB-32186] (25) Britton, E. 2668 (7) Britton, N. 493, 1241, 2076, 4200, 7611 (18) Broadway, W. 7256 (18) Bromley, G. H-180/59 (20) Brotto, M. 872 (25) Brummitt, R. 15922 (27) Bryan, G. 53 (14); 541 (52) Buchanan, J. 472, 1462, s.n. [P-00605521] (27) Buchtien, O. 414, 415, 886, 2635, 2726, 2736 (27); 3464, 3465 (36); 30, 5, 5167 (12); 5168 (21); 5172 (27); 5, 5177 (36); 00, 0 (12) Bües, C. 1737 (3) Burchell, W. 5152 (27) Burger, W. 5335 (9); 6017 (24); 6726 (21) Burrows, J. 2352 (27) Bye, R. 26011 (37) Cabrera, J. 57111, s.n. [HAJB-HFC57111] (7) Cadorin, T. 243, 1170, 1258 (25); 3164 (39) Calatayud, G. 1090, 3117, 3920 (12) Callejas, R. 2988 (21); 6008 (44) Caluff, M. 1238 (20); s.n. [BSC-MGC1129], s.n. [BSC-MGC1836], s.n. [BSC-MGC2139] (7); s.n. [BSC-MGC2442] (20); s.n. [BSCMGC2691], s.n. [BSC-MGC2768], s.n. [BSC-MGC3074], s.n. [BSC-MGC3152], s.n. [BSC-MGC3224], s.n. [BSC-MGC3270], s.n. [BSC-MGC3319], s.n. [BSC-MGC3697], s.n. [BSC-MGC4167], s.n. [BSC-MGC4190], s.n. [BSC-MGC4632], s.n. [BSC-MGC4656], s.n. A NOMENCLATOR FOR ELAPhoGLoSSUM SECTION PoLYTrIChIA Phytotaxa 535 (1) © 2022 Magnolia Press • 5 [BSC-MGC4659], s.n. [BSC-MGC4686], s.n. [BSC-MGC4687], s.n. [BSC-MGC4692], s.n. [BSC-MGC4966], s.n. [BSC-MGC5030], s.n. [BSC-MGC5560], s.n. [BSC-MGC5580], s.n. [BSC-MGC590], s.n. [BSC-MGC5939], s.n. [BSC-MGC5975], s.n. [BSC-MGC5995], s.n. [BSC-MGC5998], s.n. [BSC-MGC5999], s.n. [BSC-MGC6026], s.n. [BSC-MGC6048], s.n. [BSC-MGC6049], s.n. [BSC-MGC6050], s.n. [BSC-MGC6132], s.n. [BSC-MGC6134] (7); s.n. [BSC-MGC867], s.n. [BSC-MGC898] (20); s.n. [HAJB-MGC3319] (7); s.n. [MGC4595], s.n. [MGC-6081] (23); s.n. [BSC-MGC6003] (43) Calzada, J. 572 (24) Campos Porto, P. 2244 (42); 2636 (39) Campos, A. 499 (24) Campos, J. 4504 (52) Canela, M. 2101 (18) Canestraro, B. 123, 190 (25) Cárdenas, D. 632, 775 (10); 6191 (44) Cardona, F. 1596 (21) Carhart, M. s.n. [NY-02008198] (49) Carlson, M. 2324 (24) Carmichael, D. s.n. [BM-000] (27) Carretero, A. 872 (12) Carriker, M. 57 (27) Cascante, A. 849 (9); 1124 (18); 1722 (9) Castillón, L. s.n. [LIL-42405], s.n. [P-01380579] (27) Caxambu, M. 2112 (25) Cayola, L. 379, 2458, 2540 (12) Cerón, C. 218, 645, 1099, 3197, 4108 (44); 18618 (52); 20717 (44) Chacón, A. 160 (21); 478 (17); 1490 (21); 1895 (18) Chambi, B. 367, 369 (27); 418 (44); 420 (21) Charter, J. 36960 (27) Chase, A. 6213 (7); 6470-a (18) Chase, N. 3177, 5263, 6666, 7330 (27) Chavarría, M. 567 (17) Chickering, A. 109-a (30) Christenhusz, M. 3118 (18); 3144 (7); 4125 (18); 4706 (25); 4812 (21); 4923 (46); 4989 (25) Christophersen, E. 1148, 2362 (27) Chrysler, M. 1749 (49) Churchill, H. 4607 (24); 5778, 5779, 5780 (48) Churchill, S. 18194 (12) Clark, J. 160 (30); 206 (18); 225 (30); 650 (18); 662 (30); 875, 2621 (18); 4297 (21); 4307 (18); 4807, 4867 (30); 5204, 5211, 5246, 5289, 5299 (12); 5975 (21); 6392 (12); 6581 (24); 6596 (36); 6607 (12); 6689 (21); 6725, 6950 (12); 7802 (21); 8969 (50); 9075, 9155, 9349 (12); 9507 (44); 9652 (18); 9917 (12); 9938, 10223 (44); 10513 (20); 10761 (21) Clarke, H. 5519 (21); 8934 (18); 8985, 9241 (21) Clement, Bro. 398 (18); 711, 731 (20); 944 (7); 948 (20); 983 (18); 1247 p.p. (7); 1247 p.p. (20); 5078 (7); 6471 (24); s.n. [HAC-CSC948] (20) Clute, W. 176 (49); 309 (18) Cogollo, A. 7628 (21); 7817 (52) Colín, R. 17718 (49) Commerson, P. s.n. [E-0005, P-JU-000], s.n. [P-00605462] (27) Conant, D. 879, 1001 (44) Condack, J. 337 (42); 446 (21); 519 (27); 606 (42); 623 (27) Congdon, T. 174 (27) Conzatti, C. 799 (24) Cook, O. 90, 132 (24); 159 (21); 849, 852 (12) Cooper, J. s.n. [US-03339923] (24) Copeland, E. 158 (31); s.n. [NY-01044212], s.n. [NY-02008110] (18) Copley, H. B6495 (27) Cordeiro, I. 727 (25) 5 • Phytotaxa 535 (1) © 2022 Magnolia Press MATOS ET AL. Cornejo, X. 8066, 8074 (52) Correa, M. 186 (30); 187, 192 (10); 378 (18) Correa, M. 1885 (6); 1886 (10); 8214 (30); 11516 (10) Cortés, S. 3717 (24) Costa, M. 239, 526 (44) Cours, G. 203, 4334 (27) Cremers, G. 6108, 7464, 7947, 8945 (44) Croat, T. 863 (32); 0 (6, currently lost); 21255 (44); 22255 (22); 5 (1); 24313 (30); 27249 (10); 29488 (27); 35237 (22); 35825 (24); 36115 (21); 36664 (24); 37193 (22); 37811, 37844, 37911 (21); 38046 (30); 43068, 43070, 43134 (31); 46367, 46626 (24); 47029 (21); 51337 (12); 51343 (29); 51504 (27); 54913 (12); 55444 (52); 59970 (48); 66766 (22); 67910, 68232 (21); 69111 (31); 71354 (12); 71535 (24); 72088, 72100 (52); 88817, 88848 (12); 91093 (21); 96748 (12); 96815 (21); 96816, 96856, 97183 (12) Crosby, G. 95 (18) Crosby, M. 1007 (18); 1013 (23); 2546 (9); 2599 (32); 2722, 2724 (9); 2751 (18) Cruz, G. C61 (49) Cuadros, H. 3909 (21) Cuatrecasas, J. 369 (12); 18440 (52); 18754 (24) D’Aoust, V. s.n. [P-01405589] (34) da Costa, I. 355 (36) Dale, E. 52 (18) Dalström, S. 2214 (21) Daly, D. 8948 (44) Damasceno, E. 247 (25); 395 (42) Damazio, L. 230, 710, 730 (21); 1024 (27); 1879, 1931 (25); s.n. [Herb. no. 36797] (21); s.n. [NY-00891472] (27); s.n. [OUPR-10149], s.n. [OUPR-10151] (21); s.n. [OUPR-10754] (25); s.n. [OUPR-10757] (27); s.n. [OUPR-5911] (21); s.n. [P-01405603] (27); s.n. [P01466052] (21); s.n. [RB-01075046] (27) Daubenberger, Père 45, 46, s.n. [Filices Africae orient. germ., no. 54], s.n. [Filices Africae orient. germ., no. 60], s.n. [K-000825210] (27) Davidse, G. 24334 (6); 28731, 28795, 28940 (48); 29104, 35376 (24); 35905, 36497 (30); 36729 (22); 36794 (24) Day, E. s.n. [NY-02008112], s.n. [NY-02008123] (18) de la Sota, E. 4413 (27); 5167 (22) de Thiouville s.n. [P-0] (24) Decary, R. 14311, 16509, 17502 (27) Delgado, R. 67 (32) Díaz-Barriga, H. 4674 (49) Díaz, C. 3292 (35) Díaz, H. 6557 (34) Dickson, J. 23, 17/66 (27) Dietrich, H. 67095 (21) Dittrich, V. 379, 435, 452, 526 (25) Dodson, C. 7246, 9143, 10346 (30); 12213 (52) Doidge, E. 2482 (27) Dombrowski, L. 2887, 4496, 7481, 11390 (25) Dorante, J. 751 (24) Dorr, L. 3513 (27); 7264, 7574 (24); 7840 (38); 8565, 8875, 9066 (24); 9223 (12) Dressler, R. 1706 (24) Dudley, T. 13008 (21) Duek, J. 81 (7); E13705 (21) Dunaiski, A. 2266 (25) Dusén, P. 669 (27); 778, 2158, 3793, 7492, 8485 (25); 10619 (39); 10620 (27); 13516, 14312, 14408, 14409, 15203 (25); 18102 (21); s.n. [S-11-6714] (25) Duss, A. 185, 836, 1618 (18); 1621 (7); 4127 (4); 4148 (18); 4149 (24); 4152 (21); 4321 (7); 4458 (24); 4577 (7); s.n. [US-03339879] (21) Dutra, J. 3 (27) Dwyer, J. 8922 (1) A NOMENCLATOR FOR ELAPhoGLoSSUM SECTION PoLYTrIChIA Phytotaxa 535 (1) © 2022 Magnolia Press • 53 Eberhardt, D. 521 (28) Edwall, G. 4690 (25); s.n. [Comm. Geogr. Geol. de São Paulo, no. 4691] (21); s.n. [SP-21174] (25) Eggeling, W. 1018 (27) Eggers, H. 245 (18); 2745 (7); 2760 (18); 4997 (20); 5168, 5328 (43); 2745-b (7); s.n. [Herb. Fil. Carl Christensen, n. 309], s.n. [P01409050] (18) Eiten, G. 2082 (25) Ekman, E. 2102, 2987 (7); 3489 (30); 3880 (18); 4729 (7); 4776 (21); 5093 (20); 6981 (18); 7024 (20); 8448, 8853 (24); 9217, 10259 (43); 11562 (7); 12724 (4); 12857 (7); 12968 (18); 13405 (24); 13406 (30); 14416 (7); 11819-a (24); H-2542, H-3729 (18); H10012 (49); H12698 (22); H12816, H13572, H1402, H1866 (49); H7375 (22); H7680 (49) Engel, A. 0 (38) Engelmann, R. 172 (25); 1000, 1775 (46) Engels, M. 686, 1760, 2348 (25) Ernst, W. 1739 (24) Eugenio, Bro. s.n. [US-01566765] (27) Evans, A. 38 (9); 40 (48); 85, 223 (17); 3105 (21); 3132 (22) Ewan, J. 15663 (4); 16516 (12) Eyerdam, W. 349 (18) Faden, R. 69/526, 71/77, 72/350, 74/1340 (27) Fanshawe, D. F2430 (44) Farney, C. 2485 (46) Farrera, M. 1083 (22) Fay, A. 839 (18); 2034 (3); 2223, 2225, 3750 (12); 3839 (52) Fendler, A. 264 (24); 3, 284 (4) Fernandez, A. 256 (24); 872 (9); 4943 (44); 5339, 5531 (21); 5611 (19) Fernández, E. 2390 (12); 2822 (29); 3144 (12); 3205 (27) Fernandez, Y. 164 (41) Ferrari, F. 7 (25) Feuerer, T. 19509-a (47) Feuillet, C. 10349 (44) Fiaschi, P. 1788 (21); 3050 (46); 3243, 3250 (41); 3829, 3889, 3896 (27); 4533 (25) Figueiredo, J. 80 (27); 421, 491 (36) Figueiroa, A. s.n. [UC-723381] (24) Filho, A. 2020 (25) Fisher, J. D65-23 (18) Fletes, E. 366 (30) Flores, O. s.n. [OUPR-10760] (27) Folsom, J. 3576 (10); 3981 (22); 4266, 4526 (21) Fonseca, R. 3612 (49) Forbes, J. 155 (46) Fortes, A. 196 (21) Förther, H. 10146, 11103 (34) Forzza, R. 1209 (27) Foster, M. 685, 2288 (21); 79-94, 85-136 (12); 85-3, 85-32, 85-60 (52) Foster, R. 4159 (18); 10076, 10637, 10693 (44); 10849, 10889 (12); 13615 (18) Fournier, L. 351 (2) Franco, P. 3585 (21) Fraser-Jenkins, C. 11320 (27) Fredholm, A. 3218 (7) Fricke, R. s.n. [US-03339729] (2) Friis, I. 4087 (27) Fuentes, A. 4042 (44); 4702 (12); 4715, 4725 (29); 4843, 5938 (12); 7483 (39); 8921 (12); 9269 (36); 9326 (12); 10214 (27); 10560 (47); 11001 (12); 14741 (27); 14883 (29) Fuertes, M. 1212 (18) Galeotti, H. (24) 5 • Phytotaxa 535 (1) © 2022 Magnolia Press MATOS ET AL. Gallardo, G. 349 (30) Gamboa, B. 204 (30); 371 (17) García-Ruiz, I. 3069, 4371 (37) Garcia, A. 59 (24) García, B. 388-a (18); 388-b, 484 (24); 571, 590 (18); 666 (24); 793 (37) García, E. 4453 (12); 4494 (39) García, R. 1179 (18) Gardner, G. 55 (27) Gardner, H. 1276 (27) Garnier 290, 291 (7) Gasper, A. 599 (39); 844 (25); 2958 (51) Gastony, G. 112 (18); 375, 446, 470 (49); 703 (18) Gatti, A. 89, 255, 344 (25) Gaudichaud, C. s.n. [B-200071871] (46) Gay, H. 1551 (17) Gentry, A. 13763 (21); 22977 (35); 28627 (21) Gerdes, E. 105 (25) Gilbert, B. 5 (18); 205 (23) Gilbert, M. 214, 4190, 6319 (27) Gittins, B. 4157 (24) Glaziou, A. 950 (46); 05 (25); 3 (39); 3 (46); 3320, 3321, 4363, 4366 (27); 5366 (25); 5366 (27); 53 (42); 5368 (27); 12281 (46); 15712 (21); s.n. [P-01359824] (46); s.n. [P-01380585] (42); s.n. [P-01466051] (21); s.n. [P-01466198] (25); s.n. [US-01567510] (42) Godoy, J. s.n. [OUPR-10791] (27) Goebel, K. s.n. 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G377 (44) Gutiérrez, J. 1133, 1281 (12) Haber, W. 8585 (9) Hagemann, W. 1508 (19) Hager, J. 347 (18); 600 (49) Hahn, L. 1490 (18); s.n. [P-0055, P-0055] (34) Hall, H. 127, 161, 193, 207 (27) Hall, J. s.n. [K-000825213] (27) Hallberg, B. 1461 (24) Hamilton, C. 3944 (22) A NOMENCLATOR FOR ELAPhoGLoSSUM SECTION PoLYTrIChIA Phytotaxa 535 (1) © 2022 Magnolia Press • 55 Hammel, B. 7502 (31); 16463 (22); 16526 (30) Hammer, G. 9 (31) Handro, O. 523, 544 (25); 673 (46); 749 (21); 1215 (25); s.n. [US-01566366] (21) Hans 10 (25) Harling, G. 17957 (52); 24424 (21) Haro-Carrión, X. 186 (30) Harris, W. 1560, 7432 (7); 7433 (23); 7444 p.p., 7501 (7); 7444 p.p. (20) Hart, J. 48 (49); 168 (20); 117 p.p. (7); 117 p.p. (20); s.n. [US-03339704] (7) Hatch, W. 86 (24) Hatschbach, G. 10189, 13596, 14458, 17808, 19489, 20307, 20727, 20915-b, 22408, 25727, 27570 (25); 36206 (3); 46065, 48792, 53731 (25) Hedberg, O. 904 (27) Hekavey, C. 14 (25) Heller, A. 1076 (18) Hellwig, R. 393 (12) Henderson, Miss. s.n. [NY-02008109] (18) Henkel, T. 156, 1295 (18); 1461 (21); 5038 (44) Hennipman, E. 6891 (22); 6791 (24) Hernández, C. CICH431, CICH879 (31) Hernández, M. 460 (12) Hernandez, R. 1461 p.p. (24) Herrera, F. 1314 (27) Herrera, G. 1203 (10); 1995 (18); 3013 (30); 3166 (22); 3182, 4008, 4393, 4479 (30); 5299 (21); 5323 (26); 5348, 5372, 5472 (9); 5570 (30); 5893 (9); 5961 (21); 7851 (40); 7953 (18); 8208 (6); 9082 (9); 1456, 7855 (24) Herrera, H. 799, 1169 (30) Herzog, T. 2315 (12) Hess, W. 216, 355 (7); 358 (24); 360, 2852 (18) Hickey, R. 544 (24) Hinton, G. 1, 1867, 15609 (37) Hioram, Bro. 248 (4); 257 (7); 2428 (43); 2549 (18); 4710 (7); 6282 (20); 6350 (18); 6353 (7); 7016 (20); 2428 (43); 4710, 7040, 9383 (7); 9384 (20); 9393 (43); s.n. [HAC-HROIG7041] (20); s.n. [HAC-HROIG7050], s.n. [HAC-NSC1147] (43) Hodge, W. 1021 (24) Holdridge, L. 2147 (18); 2198 (43); 2199 (7); 2249 (49); 5133 (18); 5145 (2) Holland, D. 90 (22) Holm-Nielsen, L. 4610 (12) Holme, H. s.n. 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[K-000500] (49) Janssen, T. 2809, 2860 (27) Jaramillo, J. 10924, 11217 (44) Jaramillo, R. 5189 (21) Jardim, J. 1133 (25) Jascone, C. 711, 791 (46) Jenman, G. (23); , 20, 22 (49); 120 (18); 1458, 5, 2158 (44); s.n. [NY-02008004] (7); s.n. [NY-02008040] (20); s.n. [NY-02008095], s.n. [NY-02008099], s.n. [NY-02008111] (18); s.n. [NY-02008159] (24); s.n. [NY-02008199] (49) Jeremie, J. 235 (18) Jermy, A. 3001 (4); 11192 (18) Jiménez-Barrios, J. 1141 (24) Jiménez, A. 790 (9) Jiménez, I. 210 (39); 695 (12); 699-b, 739, 772, 775 (29); 846, 847 (36); 872 (39); 889, 890 (29); 1101, 1121 (12); 1140 (27); 1213, 1253 (29); 1300 (39); 1320 (29); 1366 (36); 1393 (39); 1395 (29); 1518 (12); 1662 (12); 1706 (36); 1714 (29); 1788 (27); 2111 (47); 2136 (28); 2144 (29); 2153 (27); 2231 (12); 2250 (27); 2255 (36); 2311 (44); 2341, 2349, 2380, 2381 (27); 2456 (12); 2564, 2677 (3); 2821 (12); 3017 (36); 3020 (12); 4853 (27) Jiménez, J. 1321 (18) Jiménez, Q. 2308 (32); 2339 (10) Johnson, D. s.n. [MO-5499038], s.n. [MO-5499039], s.n. [NY-02008202] (49); s.n. [US-03339808] (20) Johnson, H. 18, 19 (27); 675 (24); 852 (21); 858 (18) Jones, G. 1117 (49) Jörgensen, P. 1496-a (27) Jürgens, C. 75 (27); 166, 331 (27); 751 (25); s.n. [Rosenstock, Filices Austrobrasilienses, no. 149], s.n. [Rosenstock, Filices Austrobrasilienses, no. 277] (27); s.n. [Rosenstock, Filices Austrobrasilienses, no. 328] (25) K.E.R. [Kew Expedition to Roraima] 17 (21) Kanehira, R. 151 (44) Kapelle, M. 5419 (9) Karsten, H. s.n. [B-0000] (27); s.n. [B-000, LE000050, 00005, P-0035] (12); s.n. [Herbarium Filicum Carl Christensen 2134] (38) Kellermann, B. 50 (25) Kennedy, H. 2497 (10); 3026 (18); 3138 (10) Kersten, R. 0, 1172, 1482 (25) Kessler, M. 37 (29); 3, 4320 (47); 6099, 6206 (12); 6457 (27); 6482 (28); 6504, 6880 (27); 7058 (12); 7112, 7183 (36); 7283 (27); 7304 (36); 35 (); 7755 (39); 7760, 7879, 7983 (36); 8107 (44); 8129 (27); 8670 (3); 8836 (44); 9007, 9080 (29); 9085 (39); 9093 p.p. (29); 9104 (36); 9157 (12); 9300, 9450 (27); 9501, 9772 (12); 10245 (28); 10465 (47); 10527 p.p. (27); 10527 p.p. (47); 10557, 10642 (12); 10869 (3); 11321 (27); 11323 (36); 11444 (39); 11537 (12); 11721 (27); 11767, 11918 (12); 12060 (29); 12117 (27); 12160 (29); 12413, 12476, 12929 (27); 12998 (36); 13007 (39); 13049, 13305 (36); 13359 (14); 13802 (27) Kieling-Rubio, M. 436 (25) Killip, E. 5014, 5030 (32); 5262, 5277, 5298 (9); 5322 (24); 5369 (9); 5480 (32); 6733 (12); 11130, 11624 (22); 17846 (12); 26782, 28498 (44) Kirizawa, M. 1484 (25) Klawe, W. s.n. [US-03339727] (2) Klein, R. 7646, 9260 (25) Klug, G. 3330 (3); 7689, 7721, 7735, 7752, 7776 (27) Knapp, S. 8413 (44) Knox 1681 (27) Koczicki, C. 184 (25) A NOMENCLATOR FOR ELAPhoGLoSSUM SECTION PoLYTrIChIA Phytotaxa 535 (1) © 2022 Magnolia Press • 5 Køie, M. 4590 (21) Kollmann, L. 4151-b (42); 4870-a (21); 4981 (15) Koptur, S. SK-101 (24) Korte, A. 2105 (25); 2397 (39) Kozera, C. 1232, 1233 (25) Krause, R. 65, 105 (15) Kriebel, R. 3876 (32) Krieger, L. 24099, s.n. [CESJ-24099] (21) Krömer, T. 640 (44); 1044 (3); 1238 (36); 1240 (39); 1244, 1540 (28); 1686 (12); 1710 (39); 1990, 2257, 2362, 2428, 2850, 3285, 3671 (24); THO3475, THO3951 (31) Kuhlmann, J. s.n. [RB-01075032] (46) Kunkel, G. 535 (28) Kupper, W. 402, 618, 680 (9); 1629 (32) Kvist, L. 48301 (30) L’Herminier, F. 1 (18); 1 (21); 7 (24); 13 (21); 23 (18); s.n. [MO-1867292], s.n. [NY-01186184] (21); s.n. [NY-02008108] (18); s.n. [P01359791] (7); s.n. [P-01409032] (18) Labat, J.-N. 3017, 3770 (27) Labiak, P. 231, 343 (25); 986 (46); 1005 (25); 1101 (21); 2049, 3003, 3235 (25); 4074 (21); 4281 (25); 4427, 4490, 4510 (27); 4581-a (42); 4861 (39); 4865 (25); 4968 (42); 5178 (5); 5505 (21) Labillardière s.n. [P-00605466] (27) Laegaard, S. 51159 (44) Laferrière, J. 219 (44) Langsdorff, G. s.n. [M-187], s.n. [P-01380589] (46); s.n. 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[P-01464079] (44) Letouzey, R. 14557 (27) Lévy, P. 1419 (30) Lewis, J. s.n. [MO-5483759] (7) Lewis, M. 881732 (27) Liebmann, F. s.n. [GH, K, P-00] (34); s.n. [C no. LNR-50, C no. LNR-5, P-00, US-000], s.n. [US03339863] (24) Liesner, R. 12966 (24); 17784, 18578 (21); 21405, 5 (41); 21631 (19); 21736 (21); 23560, 25620 (44) Lillo, M. 2922 (27) Lima, A. s.n. [Herb. Inst. Agron. Campinas 6008] (25) Lima, J. 24, 70 (46); 282 (21) Linares, E. 2524 (24) Lindberg, G. 53 (27) 5 • Phytotaxa 535 (1) © 2022 Magnolia Press MATOS ET AL. Lindeman, J. 694 (44) Linden, J. 5 (7); 251 (49) Lindig, A. 111 p.p (12); 251 [B-00003, BM-000, K, NY-00!, NY-00!, P-00500!, P-00500!, P005003!, P-00500!] (49) Lindman, C. A893 [Exped. I. Regnell. Filic.] (27) Linneo, I. 138 (27) Liogier, Bro. 11664 (21); 12814 (49); 13471 (24); 19517 (18); 21557, 25058 (49); 25164 (21) Lira, R. 223 (24) Lisboa, M. s.n. [OUPR-10158], s.n. 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[B-200071875] (46); s.n. [M-176], s.n. [M-177], s.n. [M-178] (27); s.n. [NY-01402651] (25) Maruffa, A. 29 (25) Matos, F. 128, 202 (25); 366 (21); 380 (25); 445 (21); 485 (25); 0, 1002, 1075 (5); 1972, 1979 (25); 2021 (27); 2025 (25); 2031 (46); 2046 (25); 2050 (42); 2057 (27); 2059, 2063 (25); 2075 (22); 2076 (32); 2082 (23); 2087 (24); 2105 (6); 2112, 2120, 2133 (9); 2134 (22); 2141, 2146 (24); 2158, 2164 (9); 2165 (24); 2166 (17); 2170 (6); 2174 (17); 2176 (9); 2180 (40); 2182 (24); 2190 (10); 2191 (25); 2196 (42); 2200 (25); 2211, 2215, 2220 (27); 2253 (46); 2440 (11); 2451 (31); 2496 (16); 2501 (50); 2502 (19); 2533 (36); 2679 (46) Matschinske, G. 48 [Rosenstock, Filices Austrobrasilienses, no. 149]; s.n. 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S803 (30) Schlieben, H. 2956, 4885 (27) Schmalz, A. 114 (25) Schmitt, J. 981 (25); 1687 (39) Schnell, R. 8027, 8051, 8360, 8645 (25) Schott, A. 85 (30) Schramm, E. 29 (30) Schreiter, R. 4364 (27) Schultes, R. 781 (20); 3526 (44) Schunke, C. s.n. [US-03339819]] (19) Schunke, J. 5495, 10164, 13751 (44) Schwacke, C. 193 (44); 11596, 14150 (21); 14583 (36); 14984 (27); 15132, s.n. [ex Herb. Schwacke 13197], s.n. [ex Herb. Schwacke 13243], s.n. [ex Herb. Schwacke 14611], s.n. [ex Herb. Schwacke 5778] (25) Schwartsburd, P. 2984 (39); 2986 (25); 3042, 3044 (36); 3073 (25); 3158, 3159 (39); 4128 (36); 4139 (25); 4165, 4192 (39) Sehnem, A. 3155 (25); 4216 (27); 5193 (25); 6468, s.n. [SP-50567] (27) Seidenschnur, C. 6, 8 (6); 11 (24); 13 (9); 14 (22); 20, 21 (24); 108, 109 (30); 131-d, 132, 135, 150, 161, 185, 200 (32); 204 (22); 218, 219, 223, 227 (24) Seidenschwarz, F. 442/1 (21) Seiler, R. 130 (33); 290, 688 (11); 876 (1) Selusniaki, M. 2208 (25) Serrano, E. 323 (9) Serrano, M. 5563 (12); 5872 (27) Shafer, J. 739 (18); 3222 (7); 3226, 3586 (18); 3653 (21); 3714, 3715 (7); 4457 (18); 8862 (43); 8874 (18) • Phytotaxa 535 (1) © 2022 Magnolia Press MATOS ET AL. Shelton, G. s.n. [BSC-MGC2819], s.n. [BSC-MGC4679] (7), s.n. [BSC-MGC4680] (7); s.n. [BSC-MGC5893] (20); s.n. [BSC-MGC5954], s.n. [BSC-MGC6060] (7) Sherring, R. 130, 357 (7); s.n. [US-00067282] (23); s.n. [US-01565255], s.n. [US-01565286] (18); s.n. [US-03339814] (20); s.n. [US03340070] (49) Sieber, F. 27 [Herb. Filic. Luersen, no. 3357], 153, 280, 3400 (27) Silva, A. 119, 389, 417, 438 (25) Silva, G. 383 (12) Silva, J. 2108, 2458, 9458, s.n. [MBM-261310] (25) Silva, S. s.n. [BHCB-42718] (25) Silveira, A. 117, 2816 (27) Sim 1592 (27) Sintenis, P. 523 (7); 1330 (18); 1338, 4277 (7); 4383 (18); 6443-b (7); s.n. [P-01409055] (18) Skog, L. 4163 (10); 5154 (12) Skutch, A. 404, 818 (49) Smith, A. 48/243 (9); H487 (6); 1413 (24); 3011 (44) Smith, A.R. 953 (24); 954 (19); 1003, 1206 (4); 1242 (24); 1250 (4); 1464, 1698 (24); 1779 (10); 1880 (18); 2136 (24); 2177 (21); 2267 (24); 2299 (18); 2306 (21); 2340, 2342, 2366 (24); 2367 (9) Smith, D. 1686 (52); 2562 (24); 2665, 3854 (44); 5087 (36); 6744 (28); 7834 (24); 8346 (44) Smith, H. 380 (7); 1065 (4); 1133 (18); 1137 (7); 2688 (30) Smith, L. 1677-a (27); 2188, 2194, 2229 (25); 2258 (46) Sodiro, L. 18 (50); 192, s.n. [36/7, US-01566043] (52); s.n. [B-200071027] (30); s.n. [B-000, BM-000335, P-0053, QPLS-0, S-R-0, SI-000, US-0003] (50); s.n. [P: “Herbier Christ, 9 Fevrier 1910”] (53); s.n. [NY-02311343] (52); s.n. [P, US-catalog n. 3: “Sodiro-Rosenstock 5 Novembre 0”] (53); [P-01407625] (30); s.n. [P-01463806], s.n. [P-01463809], s.n. [P-01463812] (49); s.n. [GH, P, US-catalog n. 2786190: “Père Mille, 5 Janvier 1911”] (53); s.n. [Q-0000213] (30); s.n. [Q-000122] (50); s.n. [P-01407623, P-01407624, US-03339997] (30); s.n. [P, UC-478151, US-catalog n. 1196791: “Sodiro-Rosenstock, 5 Novembre 1920”] (53); s.n. [UC-678152] (53) Solís, F. 399 (9) Solomon, J. 5183 (12); 9309 (29); 9311, 12953, 14908, 16391 (12); 17424 (27) Somers, P. 371 (49) Soranso, F. 111 (25) Soto, A. 1567 (24) Souza, F. 75 (25); 617, 1182 (36); 1386 (25); 1428 (36); 1441 (39); 1472 (25); 1553 (39) Souza, J. 197 (36) Souza, M. 1999, 2010 (25) Spannagel, C. , ad2 (51); 164, 164-a (Santa Catarina) (25); 164-a (Rio de Janeiro), 626 (46); s.n. [Herb. n. 21192], s.n. [Rosenstock, Filices Austrobrasilienses, no. 3], s.n. [US-01570976], [US-01570977] (51); s.n. [S-11-8612] (25) Spruce, R. 16, (44); 4639 (3); s.n. [CGE-30, K, P-00] (44) Ståhl, B. 6682 (30) Standley, P. 7679 (30); 33894 (21); 34441, 35191, 35330 (24); 35398, 35428 (49); 35571 (9); 35626, 35665, 35726 (24); 37494 (30); 38187 (9); 39876 (9); 42044, 42079 (17); 49213 (24); 51693, 51755 (21); 51766 (22); 51767, 52258 (9); 52981 (30); 61034, 61523 (11); 65472, 66241, 67451 (24); 67804, 67814, 83830, 84927 (11); 85687 (24); 85977, 85979 (11); 86029 (24); 86030 (11); 86512, 89842, 90104 (24) Stannard, B. H52136 (27) Steele, M. 108 (27) Stehlé, H. 694 (24); 2284, 3431, 4485 (18) Steinbach, J. 8675 (27); 9339 (29); 9737 (12) Stergios, B. 16116, 17422, 19077, 19229, 19721, 20065, 20194, 20202 (24) Stevens, F. 2007 (18) Stevens, W. 6533 (24); 6712 (31); 9016 (30); 11367 (21); 11539 (18); 12048 (30); 0 (24); 22475 (31) Stevenson, J. 2821 (18) Steyermark, J. 448 (18); 30014 (24); 35762, 35821, 35870, 36005, 36171 (11); 37486 (34); 41554 (30); 42609 (24); 43015 (1); 43585-a (34); 44475, 3 (30); 46848, 48041, 49794, 55263 (24); 62210-a (18); 75653 (21); 89177 (44); 91710 (4); 92384, 92727 (21); 92825, 95173 (18); 97673 (21); 103552 (24); 118107, 118172 (4); 126649 (24); 126685 (4); 130312 (41) Stimson, W. 1911 (18) Stolz, A. 918 (27) A NOMENCLATOR FOR ELAPhoGLoSSUM SECTION PoLYTrIChIA Phytotaxa 535 (1) © 2022 Magnolia Press • 5 Stolze, R. 1600 (24); 1725 (44) Stork, H. 1370 (9); 1521 (17); 2030 (24) Stübel, A. 5 (52) Sucre, B. 2824 (27) Suin, L. 380 (52) Sundue, M. 516, 597, 766, 772 (12); 847 (27); 2055 (24); 3205, 3347 (12) Svenson, H. 402 (18); 403 (2) Sylvestre, L. 510 (25); 557, 679 (46); 1916 (25) Sytsma, K. 1356 (10) Tamandaré, F. 6439 (27); 6440 (25); 6555 (42) Taylor, G. s.n. [K accession n. 659-64] (7) Tejero-Díez, D. 2157, 2512, 2877 (37) Terán, J. 264 (29) Terry, M. 1583 (21) Tessmann, G. s.n. [MBM-282593] (25) Thompson, S. 5043 (24); 7417 (49) Thulin, M. 2771 (27) Tonduz, A. 9449 (10); 12239 (24); 13328 (18); 17580 (31) Torres, J. 230 (53) Torres, R. 110 (24); 137 (49); 2709, 10294 (12) Toval, N. 293 (10) Tovar, O. 2036 (27) Travassos, O. 263 (25) Trusty, J. 120, 477 (2); 513 (18) Tryon, R. 5763 (24); 6011 (53); 6532 (27); 6602 (25); 7034 (21) Tuerckheim, H. 8351, II-1465 (30) Tuomisto, H. 6182, 7767, 11447 (44) Ule, E. 229 [Herbar. Filic. Luerssen no. 12793], 286 [Herbar. Filic. Luerssen no. 13830] (25); 3 (51); 4981 (27); 6893 (44); s.n. [P01466125] (25) Underwood, L. 267, 477 (49); 924 (7); 1278 (18); 1330 (7); 1331 (18); 1358 (20); 1589 (7); 1592, 1620 (23); 2027 (7); 2065 (23); 2067 (7); 2591 (49); 2599 (20); 2698 (7); 2717 (18); 3256 (49); 3259 (20) Urban, I. s.n. [P-01380584] (46) Uribe, L. 212 (12); 5481 (52) Utley, J. 930 (9); 2809 (24) Valdespino, I. 261 (18); 0 (26) Valdez, F. 1241, 1257 (24) Valenzuela, L. 7806, 8916 (12); 12193 (21) Valerio, M. 29 (24); A92, s.n. [CR-40792] (9) Valeur, E. 650 (24); 657 (7) van der Werff, H. 5185 (44); 5817 (38); 5 (19); (6); 6299 (24); 6303 (6); 6313 (9); 6475 (24); 6594 (26); 6645 (24); 6650, 6652 (48); 6729, 6760 (21); 6836 (24); 6906 (30); 7023 (10); 7931 (24); 10264 (44); 10345, 11164 (12); 12863, 12895 (27); 13325 (44); 15252 (27); 15496 (12); 16261 (44); 16897 (27); 18048, 18308 (44); 19284, 19353 (21); 19538, 19906-a (12); 19906-b, 19938, 20276, 21188, 21224, 21292 (27); 21399-a (29); 21399-b (36); 21423 (39); 21584 (12); 21617 (21); 21712, 22005 (12); 22709 (28); 22875 (12); 24498 (30); 24772 (12); 24772 (27); 25234, 25244, 25544 (12) Vargas, C. 11392, 16822 (12) Vargas, H. 647 (24); 5287 (52) Vasco, A. 853 (52); 869 (24) Vásquez, R. 20103, 29846 (44); 30261 (35); 30573 (39); 31162 (12); 31217, 31360 (44) Vega, I. 5812 (27) Ventura, F. 8726 (31) Venturi, S. 3256 (27) Vervloet, R. 108 (15); 475 (21); (15) Vianna, F. 576 (25) Vidal, J. 1732, 1932 (42) Villa, G. 1792 (44) • Phytotaxa 535 (1) © 2022 Magnolia Press MATOS ET AL. Villalobos, J. 300 (12) Villasana, L. 161 (37) Volkens, G. 1165 (27) Völtz, R. 1077, 1472 (25); 1793 (21) von Türckheim, H. 639 (22); 954, 955, 2482 (24); 3129 (49) Vriesendorp, C. 143 (44) Wace, N. 160, T32 (27) Wachter, T. 132 (21) Wacket, M. 36, 123, s.n. [ex Herb. Museu Paulista, no. 1399] (25); s.n. [Rosenstock, Filices Austrobrasilienses, no. 284] (21); s.n. [Rosenstock, Filices Austrobrasilienses, no. 4], s.n. [SPF-107020] (25) Wadsworth, R. 503 (18) Wagner, E. s.n. [P-01325350], s.n. [P-01466202] (25) Wagner, W. 67020 (18); 71056 (9) Walcott, G. 2821 (18) Walker, T. s.n. [K accession n. 445-58.44501] (7); T10989, T10990 (18); T7085 (4) Wanderley, M. 2047 (25) Watt, M. 3, 11 (20); 19 (21); 20 (7); 98 (18); 211 (49); 10829 (21) Webster, G. 13520 (24); 13553 (18) Wegner, C. 108 (39) Wercklé, K. 180 pro parte (6); 17117 (24); s.n. [P-00249661, P-00249662] (9); s.n. [P-01466158] (6); s.n. [P-01631383] (9) Werner, F. 2118 (36); 2645, 2767 (52); 2805, 2806 (36); 1383 (53) Wettstein, R. s.n. [P-01466057] (21) White, F. 1347 (27) Widgren, J. 1302 (25) Widmer, Y. 28 (12) Wielewski s.n. [NY-000, S--3] (51) Wilbur, R. 1912 (37); 7490 (18) Wiles, J. s.n. [BM-001027745] (7) Wilkes Explor. Exped. s.n. [US-01570981] (46) Williams, L. 17560 (34); 18068 (31); 24069 (18); 25777 (11); 26212 (33) Williams, R. 1027 (3); 1031 (12); 1381 (27) Wilson, K. 455 (18); 547 (7); 551 (23); 553, 696 (7); 741 (49) Wilson, P. 72 (18); 253 (7); 321 (7) Windisch, P. 2340, 2377 (12); 5003, 5075 (46) Windisch, R. 626 (25) Wood, J. 17066 (27) Woodson, R. 355 (11); 838 (9) Woytkowski, F. 7019 (27) Wright, C. 792 (18); 3 (43); 967 (7); 972 (18); 00 (20); s.n. [P-00245229] (7) Wurdack, J. 1858, 1989 (44) Yano, O. 18963 (27) Yatskievych, G. 81-417 (37) Young, K. 2633, 3256 (27); 3394 (12); 3468 (24); 5006 (12) Yuncker, T. 5977 (22); 6251 (24); 18542 (18) Zak, V. 3599 (12); 4302 (44) Zanoni, T. 15684, 19972 (7); 19974, 20070 (18); 20121 (49); 21314 (18); 21637, 21675, 21824, 21835 (24); 22559 (18); 22608 (24); 22695 (7); 22816, 22841, 22886 (18); 23011 (7); 23050 (18); 23221 (24); 23238 (49); 24097 (18); 24150 (7); 24214 (18); 24223 (7); 24896, 24907 (49); 25076 (7); 25080 (18); 25127 (7); 25167 (18); 25170 (22); 26516, 26534 (24); 26659 (18); 26772 (7); 27087 (49); 27138 (24); 28194 (21); 29826 (18); 29836-a (24); 29850 (7); 30207 (18); 32821 (7); 33214 (18); 33218 (22); 34439 (49); 34497 (24); 35371 (18); 35380 (7); 36997 (18); 37001 (7); 37384 (24); 37584 (49); 38129 (18); 38630 (21); 39803, 39826 (24); 39995 (49); 40055 (4); 40056 (49); 41105 (21); 42431 (18); 42771 (22); 43870 (18); 43978 (7); 44197 (21) Zárate, M. 646 (12) Zogg, E. 19/12 (27) Zuill, H. 19867 (49) Zuquim, G. 66 (44) A NOMENCLATOR FOR ELAPhoGLoSSUM SECTION PoLYTrIChIA Phytotaxa 535 (1) © 2022 Magnolia Press •

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