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. We also thank several people for their invaluable assistance in the field: André Amorim, Bianca
Canestraro, Carlos Matos, Caroline Matos, Claudine Mynssen, Lana Sylvestre, Paulo Labiak, and Paulo Windisch in
Brazil; Alfredo Cascante, Caroline Matos, and Alexander Rojas in Costa Rica; Alvaro Pérez and Daniela Cevallos
in Ecuador; Alejandra Vasco, Michael Sundue, and Rafael Torres Colín in Mexico. Special thanks goes to Dr. John
Mickel for his constant encouragement and support. The drawings of Figure 4 are by Charles D. Clare, and those of
Figure 5 by Haruto Fukuda. We finally thank Marcus Lehnert, Michael Kessler, and Michael Sundue for carefully
reviewing the final version of this manuscript.
References
Alston, A.H.G. (1958) The Brazilian species of Elaphoglossum. Boletim da Sociedade Broteriana 32: 1–32.
Anderson, W.R. & Crosby, M.R. (1966) A revision of the Hawaiian species of Elaphoglossum. Brittonia 18 (4): 380–397.
https://doi.org/10.2307/2805154
Atehortúa, L. (1983) Taxonomic study of the Elaphoglossum apodum complex (Elaphoglossaceae). Ph.D. thesis, City University of New
York, New York, 214 pp.
Atehortúa, L. (1984) Contribuiciones al conocimiento de los helechos del género Elaphoglossum (Elaphoglossaceae). Parte I. Morfología
y Anatomía. Actualidades Biológicas 13: 83–93.
Atehortúa, L. (1985) Contribuiciones al conocimiento de los helechos del género Elaphoglossum (Elaphoglossaceae). Parte II. Química,
Ecología y Cladística. Actualidades Biológicas 14: 122–131.
Baker, J.G. (1872) New garden plants. Garden Chronicles 1872: 1555.
Bell, P.R. (1951) Studies in the genus Elaphoglossum III. Anatomy of the rhizome and frond. Annals of Botany (oxford) 15 (59): 347–
357.
https://doi.org/10.1093/oxfordjournals.aob.a083286
Bell, P.R. (1955) Studies in the genus Elaphoglossum IV. The morphological series in the genus and their phylogenetic interpretation.
Annals of Botany (oxford) 19 (74): 173–199.
https://doi.org/10.1093/oxfordjournals.aob.a083423
Bell, P.R. (1956) Studies in the genus Elaphoglossum IV. The morphological series in the genus and their phylogenetic interpretation, part
II. Annals of Botany (oxford) 20 (77): 69–88.
https://doi.org/10.1093/oxfordjournals.aob.a083516
Benzing, D.H., Henderson, K., Kessel, B. & Sulak, J. (1976) The absorptive capacities of bromeliad trichomes. American Journal of
Botany 63: 1009–1014.
https://doi.org/10.1002/j.1537-2197.1976.tb13183.x
Brade, A.C. (1961) O gênero Elaphoglossum (Polypodiaceae) no Brasil. I. Chaves para determinar as espécies brasileiras. rodriguésia
35–36: 21–48.
Choo, T.Y.S., Matos, F.B. & Moran, R.C. (2014) The gametophytes and young sporophytes of Elaphoglossum decursivum (Dryopteridaceae)
in Costa Rica. American Fern Journal 104: 49–57.
https://doi.org/10.1640/0002-8444-104.2.49
Christ, H. (1899) Monographie des Genus Elaphoglossum. Neue Denkschriften der Allgemeinen Schweizerischen Gesellschaft für die
Gesammten Naturwissenschaften 36: 1–159.
Christ, H. (1909) Primitiae florae Costaricencis, Filices VI. Bulletin de la Société Botanique de Genève, sér. 2, 1: 216–236.
Christensen, C. (1906) Index Filicum. H. Hagerup, Copenhagen, 744 pp.
A NOMENCLATOR FOR ELAPhoGLoSSUM SECTION PoLYTrIChIA Phytotaxa 535 (1) © 2022 Magnolia Press • 3
Christensen, C. (1932) The pteridophyta of Madagascar. Dansk Botanisk Arkiv 7: 1–253, t. 1–80.
Copeland, E.B. (1947) Genera Filicum. Chronica Botanica, Waltham, 247 pp.
Cremers, G. & Aupic, C. (2008) Typifications dans l’herbier Tournefort pour des taxons représentés par des planches iconographiques de
Plumier. Adansonia, ser. 3, 30 (1): 17–30.
Diels, L. (1902) Polypodiaceae. In: Engler, A. & Prantl, K. (Eds.) Die Natürlichen Pflanzenfamilien. Verlag von Wilhelm Engelmann,
Leipzig, pp. 139–339.
Domin, K. (1929) The pteridophyta of the island of Dominica, with notes on various ferns from tropical America. Dr. Ed. Grégr. & Son,
Prague, 259 pp.
Eaton, D.C. (1860) Filices Wrightianae et Fendlerianae, nempe Wrightianae Cubenses et Fendlerianae Venezuelanae (nonnullis
Panamensibus, etc., ex coll. A. Schott et S. Hayes interjectis) enumeratae novaeque descriptae. Bachelor of Science thesis, Harvard
University, USA.
Eaton, D.C. (1861) Filices Wrightianae et Fendlerianae, nempe in Insula Cuba a Carolo Wright et in Venezuela ab Aug. Fendler, ann.
1854–60, (nonnullis Panamensibus etc. interjectis), Enumeratae cura. Memoirs of the American Academy of Arts and Sciences, n.s.
8: 193–218.
https://doi.org/10.2307/25057969
Fée, A.L.A. (1845) histoire des Acrostichées. Mémoires sur les Familles des Fougères 2. Veuve Berger-Levrault, Strasbourg, 114 pp., t.
1–64.
Fée, A.L.A. (1852) Genera Filicum. Mémoires sur les Familles des Fougères 5. Veuve Berger-Levrault, Paris and Strasbourg, 387 pp., t.
1–30.
Fuwu, X., Faguo, W. & Mickel, J.T. (2013) Elaphoglossum. In: Wu, Z., Raven, P.H. & Hong, D. (Eds.) Flora of China, Vol. 2–3
(Pteridophytes). Science Press, Beijing; Missouri Botanical Garden Press, Saint Louis, pp. 720–723.
Gómez, L.D. & Laurito, J.G. (1986) Plantae mesoamericanae novae XI. Phytologia 60: 73–76.
https://doi.org/10.5962/bhl.part.3789
Harrison, J. (2018) Elaphoglossum backhouseanum: a real basket case! Fiddlehead Forum 45: 52–55.
Hieronymus, G. (1921) Kleine Mitteilungen über Pteridophyten III. hedwigia 62: 12–37.
https://doi.org/10.1515/islm.1921.12.1-2.98
Holttum, R.E. (1978) Lomariopsis group. In: van Steenis, G.J. & Holttum, R.E. (Eds.) Flora Malesiana, series 2: Pteridophyta, ferns and
fern allies, volume 1, part 4. National Herbarium of the Netherlands, Leiden, pp. 255–330.
Howard, R.A. (1988) Charles Wright in Cuba 1856–1867. Chadwyck-Healey, Cambridge and Alexandria, 90 pp.
Jarvis, C. (2007) order out of chaos, Linnean plant names and their types. Natural History Museum and Linnean Society of London,
London, 1016 pp.
John, S.P. & Hasenstein, K.H. (2017) The role of peltate scales in desiccation tolerance of Pleopeltis polypodioides. Planta 245: 207–
220.
https://doi.org/10.1007/s00425-016-2631-2
Karsten, H. (1869) Florae Columbiae, terrarumque adjacentium, specimina selecta, tomi II, fasciculus quintus. Dümmler, Berlin, pp.
155–188.
Kessler, M., Moran, R.C., Mickel, J.T., Matos, F.B. & Smith, A.R. (2018) Prodromus of a fern flora for Bolivia. XXXV. Dryopteridaceae.
Phytotaxa 353: 1–114.
https://doi.org/10.11646/phytotaxa.353.1.1
León, B. (2006) Lomariopsidaceae endémicas del Perú. revista Peruana de Biología 13: 906–910.
https://doi.org/10.15381/rpb.v13i2.1967
Lewis Smith, R.I. (2014) A fern cultured from Antarctic glacier detritus. Antarctic Science 26: 341–344.
https://doi.org/10.1017/S0954102013000606
Linnaeus, C. (1753) Species Plantarum, tomus II. Impensis Laurentii Salvii, Stockholm, pp. 561–1200.
Lorence, D.H. (1984) Hybridization in Elaphoglossum in the Mascarene Islands. Fern Gazette 12: 341–350.
Lorence, D.H. & Rouhan, G. (2004) A revision of the mascarene species of Elaphoglossum (Elaphoglossaceae). Annals of the Missouri
Botanical Garden 91: 536–565.
Lóriga, J., Vasco, A., Regalado, L., Heinrichs, J. & Moran, R.C. (2014). Phylogeny and classification of the Cuban species of Elaphoglossum
(Dryopteridaceae), with the description of Elaphoglossum sect. Wrightiana sect. nov. Plant Systematics and Evolution 300: 937–
951.
https://doi.org/10.1007/s00606-013-0933-4
Mabberley, D.J. (2008) Mabberley’s plant-book: a portable dictionary of plants, their classifications and uses, 3rd edition. Cambridge
University Press, New York, 1021 pp.
Manton, I. & Vida, G. (1968) Cytology of the fern flora of Tristan da Cunha. Proceedings of the royal Society of London, series B,
• Phytotaxa 535 (1) © 2022 Magnolia Press
MATOS ET AL.
Biological Sciences 170 (1021): 361–379.
https://doi.org/10.1098/rspb.1968.0045
Matos, F.B. (2014) Typification of some species names in Elaphoglossum section Polytrichia (Dryopteridaceae) from Brazil. Acta Botanica
Brasilica 28: 404–407.
https://doi.org/10.1590/0102-33062014abb3389
Matos, F.B. (2015) Systematic studies of Elaphoglossum section Polytrichia (Dryopteridaceae). Ph.D. thesis, City University of New
York, New York, 243 pp.
Matos, F.B. (2020a) Elaphoglossum in Flora do Brasil 2020. Jardim Botânico do Rio de Janeiro. Available from: http://floradobrasil.jbrj.
gov.br/reflora/floradobrasil/FB91002 (accessed: 2 July 2021).
Matos, F.B. (2020b) (2767) Proposal to reject the name Acrostichum webbii (Elaphoglossum webbii) (Dryopteridaceae). Taxon 69: 1108–
1109.
https://doi.org/10.1002/tax.12332
Matos, F.B. & Mickel, J.T. (2014) The Brazilian species of Elaphoglossum sect. Polytrichia (Dryopteridaceae). Brittonia 66: 371–395.
https://doi.org/10.1007/s12228-014-9352-2
Matos, F.B. & Moran, R.C. (2013) Elaphoglossum clathratum sp. nov. (Dryopteridaceae) from the eastern side of the Andes in Ecuador.
Nordic Journal of Botany 31: 442–445.
https://doi.org/10.1111/j.1756-1051.2012.00039.x
Matos, F.B. & Moran, R.C. (2017) Elaphoglossum mickeliorum (Dryopteridaceae), a new species of Elaphoglossum sect. Polytrichia from
Peru. Brittonia 69: 91–95.
https://doi.org/10.1007/s12228-016-9450-4
Matos, F.B. & Vasco, A. (2015) Elaphoglossum fendleri (Dryopteridaceae), a new species of Elaphoglossum sect. Lepidoglossa from
Venezuela, and the identity of Elaphoglossum ornatum. Brittonia 67: 144–149.
https://doi.org/10.1007/s12228-015-9371-7
Matos, F.B., Prado, J. & Moran, R.C. (2015) (042–043) Proposal to modify Article 9 Note 1 and the Glossary of the Melbourne Code.
Taxon 64: 649.
https://doi.org/10.12705/643.26
Matos, F.B., Vasco, A. & Moran, R.C. (2018) Elaphoglossum doanense and Elaphoglossum tonduzii: new members of Elaphoglossum
sect. Squamipedia (Dryopteridaceae) and their significance for inferring the evolution of rhizome habit and nest-forming leaves
within the genus. International Journal of Plant Sciences 179: 296–313.
https://doi.org/10.1086/696687
Matos, F.B., Labiak, P.H. & Moran, R.C. (2019) The Decursivum Group of Elaphoglossum sect. Polytrichia (Dryopteridaceae): nonsubulate species nested in an otherwise subulate-scaled clade. Annals of the Missouri Botanical Garden 104: 400–428.
https://doi.org/10.3417/2019371
Matos, F.B., Lóriga, J. & Moran, R.C. (2021) Monograph of Elaphoglossum sect. Polytrichia subsect. Apoda (Dryopteridaceae). Systematic
Botany 46: 764–789.
https://doi.org/10.1600/036364421X16312067913462
Mettenius, G.H. (1856) Filices horti Botanici Lipsiensis. Verlag von Leopold Voss, Leipzig, 135 pp., t. 1–30.
Mettenius, G.H. (1864) Prodromus Florae Novo-Granatensis, Filices. Annales des Sciences Naturelles, Botanique, sér. 5, 2: 193–271.
Mickel, J.T. (1980) New Elaphoglossums (Elaphoglossaceae) from Mexico. Brittonia 32: 334–339.
https://doi.org/10.2307/2806728
Mickel, J.T. (1981) Elaphoglossum. In: Stolze, R.G. (Ed.) Ferns and fern allies of Guatemala, Part II, Polypodiaceae. Fieldiana, Botany,
new series, 6: 210–238.
Mickel, J.T. (1985) Elaphoglossum. In: Smith, A.R. (Ed.) Pteridophytes of Venezuela, an annotated list. Published by the editor, pp.
78–107.
Mickel, J.T. (1987) New species of Elaphoglossum (Elaphoglossaceae) from northern South America. Brittonia 39: 313–339.
https://doi.org/10.2307/2807127
Mickel, J.T. (1991) Elaphoglossum. In: Tryon, R.M. & Stolze, R.G. (Eds.) Pteridophyta of Peru, Part IV, 17: Dryopteridaceae. Fieldiana,
Botany, new series 27: 111–166.
Mickel, J.T. (1992) New species of the fern genus Elaphoglossum from Mesoamerica. Novon 2: 368–382.
https://doi.org/10.2307/3391496
Mickel, J.T. (1995a) Elaphoglossum. In: Moran, R.C., Riba, R., Davidse, G., Sousa, M. & Knapp, S. (Eds.) Flora Mesoamericana,
volumen 1: Psilotaceae a Salviniaceae. Universidad Nacional Autónoma de México, Mexico City, pp. 250–283
Mickel, J.T. (1995b) Elaphoglossum. In: Steyermark, J.A., Berry, P.E. & Holst, B.K. (Eds.) Flora of the Venezuelan Guayana, vol. 2:
Pteridophytes, Spermatophytes p.p. Missouri Botanical Garden Press, Saint Louis, pp. 89–105.
A NOMENCLATOR FOR ELAPhoGLoSSUM SECTION PoLYTrIChIA Phytotaxa 535 (1) © 2022 Magnolia Press • 5
Mickel, J.T. & Atehortúa, L. (1980) Subdivision of the genus Elaphoglossum. American Fern Journal 70: 47–68.
https://doi.org/10.2307/1546218
Mickel, J.T. & Beitel, J.M. (1988) Pteridophyte flora of Oaxaca, Mexico. Memoirs of the New York Botanical Garden 46: 1–568.
Mickel, J.T. & Smith, A.R. (2004) The pteridophytes of Mexico. Memoirs of the New York Botanical Garden 88: 1–1055.
Moore, T. (1857–1862) Index Filicum: a synopsis, with characters, of the genera, and an enumartion of the species of ferns, with synonyms,
references, etc. William Pamplin, London, 396 pp.
https://doi.org/10.5962/bhl.title.55301
Moore, T. (1882) New garden ferns. The Gardners’ Chronicles, n.s., 17: 672–673.
Moran, R.C., Klimas, S. & Carlsen, M. (2003) Low-trunk epiphytic ferns on tree ferns versus angiosperms in Costa Rica. Biotropica 35:
48–56.
https://doi.org/10.1111/j.1744-7429.2003.tb00261.x
Moran, R.C. & Mickel, J.T. (2004) Three new neotropical species of Elaphoglossum (Elaphoglossaceae) with subulate scales. Brittonia
56: 200–204.
https://doi.org/10.1663/0007-196X(2004)056[0200:TNNSOE]2.0.CO;2
Moran, R.C., Hanks, J.G. & Rouhan, G. (2007) Spore morphology in relation to phylogeny in the fern genus Elaphoglossum
(Dryopteridaceae). International Journal of Plant Sciences 168: 905–929.
https://doi.org/10.1086/518269
Moran, R.C., Labiak, P.H. & Sundue, M. (2010a) Phylogeny and character evolution in bolbitidoid ferns (Dryopteridaceae). International
Journal of Plant Sciences 171: 547–559.
https://doi.org/10.1086/652191
Moran, R.C., Labiak, P.H. & Sundue, M. (2010b) Synopsis of Mickelia, a newly recognized genus of bolbitidoid ferns (Dryopteridaceae).
International Journal of Plant Sciences 171: 547–559.
https://doi.org/10.1086/652191
Morton, C.V. (1968) The fern collections in some European herbaria. American Fern Journal 58: 158–168.
https://doi.org/10.2307/1545951
Petiver, J. (1712) Pteri-graphia Americana, icones continens plusquam 400 filicum variarum specierum, etc. Publisher not identified,
London, 23 pp.
Pichi Sermolli, R.E.G. & Bizzarri, M.P. (2005) A revision of Raddi’s pteridological collection from Brazil (1817–1818). Webbia 60:
1–393.
https://doi.org/10.1080/00837792.2005.10739597
Plumier, C. (1705) Traité des fougères de l’Amerique. Imprimerie Royale, Paris, 146 pp., t. 1–170.
Presl, C.B. (1836) Tentamen Pteridographie. Typis Filiorum Theophili Haase, Prague, 290 pp., t. 1–12.
Presl, C.B. (1851) Epimeliae Botanicae. Abhandlungen der Königlichen Böhmischen Gesellschaft der Wissenschaften, ser. 5, 6: 361–624,
t. 1–15.
Proctor, G.R. (1977) Pteridophyta. In: Howard, R.A. (Ed.) Flora of Lesser Antilles, Leeward and Windward Islands, vol. 2. Harvard
University, Cambridge, 414 pp.
Proctor, G.R. (1985) Ferns of Jamaica. British Museum (Natural History), London, 631 pp.
Proctor, G.R. (1989) Ferns of Puerto Rico and the Virgin Islands. Memoirs of The New York Botanical Garden 53: 1–389.
QGIS Development Team (2013) Quantum GIS, version 2.0. Geospatial Foundation, Vancouver. Available from: http://qgis.org/ (accessed:
2 July 2020)
Racine, J.S. (2012) RStudio: A platform-independent IDE for R and Sweave. Journal of Applied Econometrics 27: 167–172.
https://doi.org/10.1002/jae.1278
Raddi, G. (1825) Plantarum brasiliensium nova genera et species novae vel minus cognitae collegit et descripsit Iosephus raddius. Pars
I. (Filices). Typographia Aloisii Pezzati, Florence, 101 pp., t. 1–84.
Rojas-Alvarado, A.F. (1996a) Twelve new species of Elaphoglossum (Elaphoglossaceae) from Costa Rica and Panama. Brenesia 45–46:
7–26.
Rojas-Alvarado, A.F. (1996b) Aportes a la flora pteridophyta costarricense I. Informes. Brenesia 45–46: 1–6.
Rojas-Alvarado, A.F. (2002) New species, new combinations and new distributions in neotropical species of Elaphoglossum
(Lomariopsidaceae). revista de Biología Tropical 50: 969–1006.
Rojas-Alvarado, A.F. (2003) Notes on Elaphoglossum (Lomariopsidaceae) section Polytrichia subsection hybrida in Mexico and Central
America. revista de Biología Tropical 51: 33–48.
Rojas-Alvarado, A.F. (2009) Two new species and a new combination in Elaphoglossum sect. Polytrichia subsect. Apoda (Dryopteridaceae)
from Costa Rica and Panama. Brittonia 61: 293–300.
https://doi.org/10.1007/s12228-009-9078-8
• Phytotaxa 535 (1) © 2022 Magnolia Press
MATOS ET AL.
Rojas-Alvarado, A.F. (2010) New species and new records in Elaphoglossum sect. Polytrichia subsect. hybrida (Dryopteridaceae) from
the Neotropics. American Fern Journal 100: 172–179.
https://doi.org/10.1640/0002-8444-100.3.172
Rojas-Alvarado, A.F. (2017) A new species and three hybrids in the ferns from Cocos Island, Puntarenas, Costa Rica. Acta Botanica
Malacitana 42: 91–99.
https://doi.org/10.24310/abm.v42i1.2383
Rojas-Alvarado, A.F. (2019) Corrections in recently described species of ferns and lycophytes from the neotropics. open Access Library
Journal 6: e5172.
https://doi.org/10.4236/oalib.1105172
Rouhan, G. (2020) Synoptic revision of the fern genus Elaphoglossum Schott ex J.Sm. (Dryopteridaceae) in Madagascar, with the
description of 23 new taxa, all but one endemic. PeerJ 8:e10484.
https://doi.org/10.7717/peerj.10484
Rouhan, G. & Cremers, G. (2006) Lectotypifications of some American Acrostichum and Elaphoglossum (Elaphoglossaceae). Systematics
and Geography of Plants 76: 179–183.
Rouhan, G., Dubuisson, J.-Y., Rakotondrainibe, F., Motley, T.J., Mickel, J.T., Labat, J.-N. & Moran, R.C. (2004) Molecular phylogeny of
the fern genus Elaphoglossum (Elaphoglossaceae) based on chloroplast non-coding DNA sequences: contributions of species from
the Indian Ocean area. Molecular Phylogenetics and Evolution 33: 745–763.
https://doi.org/10.1016/j.ympev.2004.08.006
Roux, J.P. (1982) The fern genus Elaphoglossum Schott (Filicales) in South Africa. Journal of South African Botany 48: 481–526.
Rovirosa, J.N. (1909) Pteridografía del sur de México. Imprenta de Ignacio Escalante, Mexico City, 298 pp., t. 1–70.
Schelpe, E.A.C.L.E. (1969) Reviews of tropical African Pteridophyta. Contributions from the Bolus herbarium 1: 1–132.
Schuettpelz, E. & Pryer, K.M. (2007) Fern phylogeny inferred from 400 leptosporangiate species and three plastid genes. Taxon 56:
1037–1050.
https://doi.org/10.2307/25065903
Shimek, B. (1897) The ferns of Nicaragua, an account of the ferns collected by the Nicaragua botanical expedition of the State University
of Iowa in 1893. Bulletin from the Laboratories of Natural history of the State University of Iowa 4: 116–224, t. 1–20.
Skog, J.E., Mickel, J.T., Moran, R.C., Volovsek, M. & Zimmer, E.A. (2004) Molecular studies of representative species in the fern genus
Elaphoglossum (Dryopteridaceae) based on cpDNA sequences rbcL, trnL-F, and rps4-trnS. International Journal of Plant Sciences
165: 1063–1075.
https://doi.org/10.1086/423877
Smith, A.R., Pryer, K.M., Schuettpelz, E., Korall, P., Schneider, H. & Wolf, P.G. (2006) A classification for extant ferns. Taxon 55:
705–731.
https://doi.org/10.2307/25065646
Smith, J. (1846) Additions to the “Hortus Kewensis”. Curtis’s Botanical Magazine 72 (Companion): 1–39.
Sodiro, A. (1893) Cryptogamae vasculares quitenses adjectis speciebus in aliis provinciis ditionis ecuadorensis hactenus detectis. Typis
Universitatis, Quito, 656 pp., t. 1–7.
https://doi.org/10.5962/bhl.title.44791
Sorsa, V. (1966) Chromosome numbers of Elaphoglossum and hymenodium. Nature 210: 334.
https://doi.org/10.1038/210334a0
Stafleu, F.T. & Cowan, R.S. (1976) Taxonomic literature: a selective guide to botanical publications and collection dates, commentaries
and types, vol. 1 (A–G). Bohn, Scheltema & Holkema, Utrecht, 1136 pp.
https://doi.org/10.5962/bhl.title.48631
Stafleu, F.T. & Cowan, R.S. (1985) Taxonomic literature: a selective guide to botanical publications and collection dates, commentaries
and types, vol. 5 (Sal–Ste). Bohn, Scheltema & Holkema, Utrecht, 1066 pp.
Thiers, B. (2021) Index herbariorum: A global directory of public herbaria and associated staff. Available from: http://sweetgum.nybg.
org/science/ih/ (constantly updated).
Tryon, A.F. (1963) Hermann Karsten, his collections and the Flora Columbiae. Taxon 12: 103–105.
https://doi.org/10.2307/1217198
Tryon, R.M. & Tryon, A.F. (1982) Ferns and allied plants with special reference to tropical America. Springer-Verlag, New York, 857
pp.
https://doi.org/10.1007/978-1-4613-8162-4
Turland, N.J., Wiersema, J.H., Barrie, F.R., Greuter, W., Hawksworth, D.L., Herendeen, P.S., Knapp, S., Kusber, W.-H., Li, D.-Z., Marhold,
K., May, T.W., McNeill, J., Monro, A.M., Prado, J., Price, M.J. & Smith, G.F. (2018) International Code of Nomenclature for algae,
fungi, and plants (Shenzhen Code). Regnum Vegetabile 159. Koeltz Botanical Books, Glashütten, 254 pp.
A NOMENCLATOR FOR ELAPhoGLoSSUM SECTION PoLYTrIChIA Phytotaxa 535 (1) © 2022 Magnolia Press •
https://doi.org/10.12705/Code.2018
Vareschi, V. (1969) Elaphoglossum. In: Lasser, T. (Ed.) Flora de Venezuela, vol. 1, tomo 2. Talleres Gráficos Universitarios, Mérida, pp.
787–866.
Wagner, W.H. (1954) A Bolivian Elaphoglossum of unique leaf structure. Bulletin of the Torrey Botanical Club 81: 61–67.
https://doi.org/10.2307/2482168
Walker, T.G. (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. [M-15633] (21)
Goldenberg, R. 985 (44)
Goldgewicht, A. 1239 (9)
Gollmer, J. s.n. [B-200069756] (4)
Gómez, L. 25 (9); 417 (18); 920, 1191, 2220, 2224 (9); 2244-b p.p., 2256 (24); 3347 (2); 4508 (18); 18042 (2); 18775 (21); 18826, 18835,
19179 (24); 19193 (18); 20180, 20209 (9); 22264 (24); 22964 (40); 23549 (24); 3 (10)
González, J. 1196, 1209 (2)
Gonzalez, S. 2175 (44)
Graf, L. 12 (18)
Graham, J. 5330 (21)
Grant, J. 01-3949, 92-01661 (24); 92-01806 (32); 92-01833, 92-01834-b (24); 92-01929 (30); 92-01942 (24)
Granville, J. 1747, 1799, 8715, 11502 (44)
Grayum, M. 3779 (22); 3860, 5087 (24); 5294 (11); 5376 (22); 7218 (24); 7277 (17); 7399 (24); 7807 (21); 9341 (18); 10831 (22); 11068
(21)
Greenway, P. 3189, 3867, 8684, 9598, 13213 (27)
Gremmen, N. T070426 (27)
Grijalva, A. 297 (22)
Grimshaw, J. 93/283, 93/781, 93/781A, 93/782, 93/789, 93/823 (27)
Grubb, P. 79 (21); P63 (12)
Guilding, L. s.n. [E-0003, K-000505, K-0005050, K-000505] (7)
Guppy, N. 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. [K-000299934] (18)
Holst, B. 3875 (30); 5204 (22); 5525, 5862 (30); 5924 (22)
Hoover, W. 1845 (21); 3356 (12)
Horich, C. 123 (24); 157 (22); 344 (9); 350, 482 (24); 58040-1 (18); 59042-1 (9)
Horwath, A. 83 (24)
Hoshizaki, B. 4287 (18)
Howard, R. 9111, 9378 (49); 15204 (18); 15549 (7); 15563 (18); 16064 (18)
Huamantupa, I. 4405 (35)
Huaylla, H. 72 (29); 352 (12); 775 (21); 1074, 1075 (29); 1415 (12); 1417 (3); 1506 (27); 1610, 1715 (12); 1846 (27); 2466 (12); 2778
(29); 2892 (12)
Humblot, M. s.n. [P-003] (27)
Hunnewell, F. 19428 (7)
Hunt, D. 6339 (25)
Husnot, P. 312 (7)
Husnot, T. 309 (18); 313 (24)
Hutchison, P. 3071 (19)
Idárraga, A. 2236 (24)
Iltis, H. 1066 (12)
im Thurn, F. 266 (21)
5 • Phytotaxa 535 (1) © 2022 Magnolia Press
MATOS ET AL.
Imray, J. 47 (18)
Ingela, G. 95-M014 (24)
Ingram, S. 1458 (9); 1544 (31); 1557 (6); 1582 (22); 1857 (9)
Jack, J. 8058 (20)
Jaimes, M. 179 (21)
Jakubowsky, G. T65, T232 (27)
James, W. 58 (6)
Jameson, W. 14, 461, s.n. [B-200072084], s.n. [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. [P-01405572] (27)
Lankester, C. 620 (21); 727 (24); 829 (40); s.n. [US-03339924] (24)
Larsen, K. 35389 (18)
Last, J. s.n. [K-000355], s.n. [K-000855544], s.n. [US-03339969] (27)
Lavanchy, M. s.n. [F-C0385314F] (27)
Lechler, W. 00 (27)
Lehmann, F. 2261 (19); 8544 (12); BT842, BT872, BT1214 (19)
Lehnert, M. 1283 (12); 1314 (53)
Leist, N. 2200 (44)
Leite, J. 2481, 3147, 3885 (27)
Lellinger, D. 106 (30); 245 (21); 405 p.p. (24); 439 (18); 476 (24); 490 (18); 571 (30); 649 (10); 821 (22); 865 (24); 1077, 1104 (6); 1218
(18); 1383 (9); 1408 (22); 1706 (24); 1799 (17); 1861 (30); 1905 (9); 1955 (6); 1966 (24)
Lemieux, T. 2262 (22)
Lent, R. 1217 (24); 2863 (9); 3539, 3584 (24)
León, Bro. 2078 (27); 2199 (12); 2400 (27); 9841 (20); 10524, s.n. [HAC-LS10527], s.n. [HAC-LS10527], s.n. [HAC-LS17198] (7); s.n.
[HAC-LS9841] (20)
Leonard, E. 9571 (18); 14594 (18)
Lepervanche-Mézières, M. s.n. [P-0055] (27)
Lépiz, E. 343 (18); 351 (2)
Leprieur. F. 38, 58, 244, 316, s.n. [P-01359790], s.n. [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. [OUPR-10158] (21)
Little, E. 9156 (52)
Lloyd, F. 318 (18); 4130, 4137 (24); 4148, 4159, 4162 (30); 4186 (21); 4187 (24); 4188 (18); 4197 (24); 4199 (22); 4205, 4216 p.p., 4221
(24); 4226, 4227 (32); 4236 (24); (32); 4255 (24)
Lobão 1869 (42)
Lock, J. 69/14 (27)
Loomis, H. s.n. [US-00835666] (18)
Lopes, M. 429 (5)
López , P. s.n. [BSC-MGC2590] (7)
López Figueiras, M. 2225 (20)
Lopez, M. 141 (18)
Lorea, F. 2225, 3380, 4482 (37)
Lorence, D. 1608, 2496, 2519 (27); 3873 (24); 4214 (49); 15024, 15635, 15823-a (27)
Lóriga, J. 22, 44, 56 (20); 68, 69, 149 (7); 155 (20); 162 (24); 163 (7); 174, 175 (20); 203, 263, 268, 280, 321, 344, 422, 430, 457 (7); 486,
488, 490, 491 (43)
Lötschert, W. 369 (34)
Loveridge, M. 299 (27)
Lovett, J. 613, 2071 (27)
Lozano, G. 4441 (24)
Lozano, R. 1709 (12)
Luer, C. 5085 (12)
Luerssen, C. 110, 2106 (27)
Luetzelburg, P. 265 (25); 2360 (27); 6864 (46); 12954, 12959, 18929 (25)
Luke, W. 11158 (27)
Lumer, C. 1321 (24)
Luna, C. 9306 (37)
Lundell, C. 16456 (30)
Luteyn, J. 8408 (50); 8549, 9014, 9112 (44); 11601 (24); 12528 (12); 13629 (36); 13728 (29)
Lutz, A. 1752 (25)
Lyonnet, E. 3, 16-b (37); 1359 (49); 2863, 3335 (37)
Macbride, J. 3373 (49); 4690 (12)
Macedo, T. 134 (25)
Madison, M. 664 (49); 735, 764 (18); 765 (30); 2516 (21); 4721 (52)
Madriñán, S. 247 (18)
Maguire, B. 27741, 28490, 28560, 29172 (21); 40570 (18)
Maia, L. 536 (41)
Malan, L. 97 (27)
Mamede, M. 323, 371 (25)
Mandon, G. 1542 (12)
Mann, G. 663, 1138, 1377 (27)
Markgraf, F. 3619 (39)
Marques, M. 294 (25)
Marquete, R. 909 (46)
Martínez, E. 19796 (33); 22907 (22); 23317 (30); 23969 (24); 25922 (30)
Martínez, K. 404 (9)
Martínez, O. 1791, 1882 (27)
A NOMENCLATOR FOR ELAPhoGLoSSUM SECTION PoLYTrIChIA Phytotaxa 535 (1) © 2022 Magnolia Press • 5
Martius, K. s.n. [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. [Rosenstock, Filices Austrobrasilienses, no. 149] (27)
Mattos-Silva, L. 4480 (25)
Matuda, E. 18795 (37)
Maublanc, A. 618 (21)
Maxon, W. 171 (7); 176 (18); 219 (24); 221 (31); 245 (24); 275 (9); 401 (21); 402 (9); 653 (49); 700 (21); 713 (20); 782 (18); 818, 882
(7); 903 (23); 904, 1188 (49); 1313 (20); 1453 (7); 1509 (49); 1526 (20); 1569 (18); 1740 (7); 1753 (18); 1827, 1914 (7); 1917 (18); 1919
(23); 2248 (7); 2249 (18); 2395 (7); 2409 (23); 2782 (20); 3233, 3287 (24); 4071, 4173 (7); 4334 (43); 4335 (7); 5035, 5036 (32); 5061
(22); 5064 (24); 5067 (22); 5239 (32); 5506 (24); 5512, 5522 (9); 5583 (6); 5629 (24); 5820 (30); 7962 (9); 8077 (24); 8184, 8348, 8443
(9); 8520, 8834 (7); 8853 (18); 8930 (7); 9058 (18); 9076 (23); 9081 (18); 9120, 9129 (23); 9138 (7); 9360 (18); 9379 (7); 9380, 9546
(23); 10545 (7); 10552 (18)
Mazé, H. 175 (24); 779, 872 (21); s.n. [P-01409018] (18)
Mazziero, F. 897 (42); 1044 (36); 1067 (25); 1121 (39)
McAlpin, B. 85-71 (32)
McConnell, F. 602 (21)
McLoughlin, A. 106 (27)
McPherson, G. 14090 (18); 12204-a (21)
McVaugh, R. 11757, 14319, 17836, 21439, 23196 (37)
Meave, J. 1723 (24)
Meier, W. 1990 (4); 2596 (21); 4276, 4626 (24); 10339 (18); 10726 (4); 10753, 10778 (21); 13389 (4); 13390 (24); 13462 (21); 17108
(18)
Mejía, F. 242 (24)
Mejía, M. 27 (18); 733, 1620 (18); 1621, 4976 (7); 7585, 7644, 7653 (49); 18249-b (7); 18330 (24); 18350 (18)
Mejland, Y. 617, 690, 813, 1148 (27)
Mélinon, M. 353, 386 (44)
Mellado, L. 1630 (36); 1709, 1959 (39); 2312 (14); 2373, 2398 (52); 2618 (21); 2836 (44); 2909, 2952 (12)
Melo, L. 137 (25); 140 (27); 193 (39); 227 (25); 234 (39); 235 (36)
Méndez, A. 8041 (24)
Mendieta, G. 578, 662 (53)
Mendoza, A. 102 (37)
Merz 269 (18)
Meyer, T. 14037 (27)
Michelon, C. 341, 562 (25)
Mickel, J. 0 (37); 1003, 1018 (31); 1083 (24); 1084 (31); 1848 (6); 2444 (24); 2657 (9); 2962 (21); 2977, 3019 (24); 3067 (22); 3098
(32); 3129 (24); 3131 (21); 3150 (22); 3418 (21); 3419 (22); 3772, 4052 (49); 4119-a (12); 4119-b (49); 4904 (12); 5229 (31); 5323, 5325
(49); 5646 (24); 5706 (31); (22); 7040 (49); 7101 (24); 7205 (22); 8095 (21); 8264, 8384, 8424 (49); 8544 (18); 8597 (7); 8753 (49);
8791 (4); 8848 (49); 8885 (24); 8936, 8952 (49); 8985 (22); 8988 (7); 8999 (18); 9065 (7); 9072 (18); 9127 (24); 9190 (18); 9207 (7);
9480 (4)
Miers, J. 16 (25); s.n. [BM-000, K-0000] (36)
Mille, A. 4 (24); 185 (49); 213 (12), s.n. [NY-01403032] (12); s.n. [P-01463810] (49); s.n. [US-01566042] (52)
Miller, G. 1722 (30)
Miller, J. 739 (18); 2195 (44)
Moeller, F. s.n. [P-01466124] (25)
Molesworth-Allen, B. 3628 (27)
Molina, A. 14654 (11)
Monro, A. 2016 (1)
Monteagudo, A. 5277 (44)
Monterrosa, J. 1480 (34)
Montgomery, J. 85-33 (6); 85-34 (9); 85-96 (11)
0 • Phytotaxa 535 (1) © 2022 Magnolia Press
MATOS ET AL.
Montoya, J. s.n. [NY-01402788] (18)
Mora, E. 950 (18); 1257 (30)
Morado, D. 35 (37)
Moraes, P. 2715, 2752 (5)
Moraga, M. 194 (1); 303 (17); 327 (48)
Morales, C. 1800 (12)
Morales, J. 883 (30); 3792 (9); 3796, 4762 (24)
Moran, R. 2446 (21); 3217 (18); 3218 (6); 3227 (21); 3236 (6); 3279, 3291, 3312 (9); 4109 (10); 5125, 5130, 5137, 5146 (12); 5299 (30);
5396 (52); 5813, 5858 (24); 6016 (52); 6059, 6153 (44); 6321 (10); 6355 (24); 6364 (17); 6379 (9); 6694, 6704 (52); 6724 (24); 6937 (30);
6950 (21); 6998 (17); 7440 (16); 7474 (44); 7532 (52); 7557 (12)
Moreno, P. 18859 (24); 19728 (31)
Mori, S. 25586 (44)
Moritz, J. 43 (24); 234 (4); 235 p.p., 324 (4)
Morley, B. 548 (7); 711 (18); 921 (7)
Morris, M. 232, 271 (30)
Morton, C. 3179, 3194 (7); 6211 (18); 7585, 7634 (31); 9507, 9577 (20)
Mosén, C. 2249, 2250 (27); 2633, 3047 (25); 3727 (21); 4640, 4641 (27)
Mostacero, J. 129, 796 (4); 826 (21); 865 (4)
Mota, R. 1566 (25); 1755, 2729 (21); 3183 (25)
Motley, T. 2912 (27)
Motta, J. 531, 994, 2412 (25)
Müller, F. 435 (31); 1802 (24)
Müller, F. s.n. [NY-02311324] (31)
Münch, G. (31)
Munn-Estrada, X. 1726, 1758 (24)
Murillo, M. 104 (24)
Mynssen, C. 1335 (42); 1336 (46)
Nadeaud, M. 80, s.n. [P-01403530] (25)
Nash, G. 1067 (7); 1067 (18)
Navarrete, H. 1542, 1579 (52); 1595 (12); 1766 (12); (52)
Navarro, E. 657, 877 (9); 929 (32)
Nee, M. 9641, 11227 (18); 24959 (24); 49391-a (12)
Neill, D. 3525 (10); 7765 (44); 11363 (18)
Nelson, C. 3794 (24)
Nevers, G. 8477 (21)
Neves, D. 749 (36)
Nicholls, H. 193 (7)
Nicolau, S. 979 (25)
Nicolson, D. 4067 (18)
Niño, M. 1549 (24)
Nonato, F. 820 (25)
Novelino, R. 798 (25); s.n. [OUPR-4019] (21)
Ntezurubanza, L. 40 (27)
Nuñez, A. 1040 (31)
Núñez, P. 8348 (12)
Núñez, R. 158 (39); 164, 229, 521 (29); 656 (12); 701, 754-b (29)
Ocampo, R. 882 (18)
Øllgaard, B. 860, 907, 982 (52); 1314 (44); 2289, 2459 (52); 2646 (21); 3000 (16); 3062 (52); 9153 (21); 9229 (12); 34926, 35254, 35312
(44); 37736 (52); 38921, 38972 (44); 90446, 98027 (12); 98312 (21); 98473 (44); 98507 (19); 100687 (30); 105577, 105587 (12); 105682
(21)
Onraedt, M. 691290 (27)
Orcutt, C. 4009 (34)
Ortega, F. 2744 (24); (38)
Ortega, R. 530 (34)
A NOMENCLATOR FOR ELAPhoGLoSSUM SECTION PoLYTrIChIA Phytotaxa 535 (1) © 2022 Magnolia Press •
Osmaston, H. 2167-a, 2190, 2482 (27)
Otero, J. 14 (18)
Oviedo-Brenes, F. JBW20 (32)
Pabón, M. 1011 (44)
Pabst, G. 4701, 4758 (46); 6979, 6986, 8901 (25)
Paciencia, M. 2141, 2158 (25)
Paget-Wilkes, C. 818 (27)
Palacios-Rios, M. 18 (24); s.n. [BSC-MGC3817], s.n. [BSC-MGC3822] (7)
Palacios, W. 394, 434 (44); 3813 (12); 4994 (52); 5985 (21); 6745 (12); 7598 (44); 8212 (12); 8397 (21); 8632 (44); 03 (8); 13630
(18); 16196 (44)
Papenfuss, G. s.n. [US-01565251] (18)
Parish, C. s.n. [P-00605468] (27)
Pedraza-Peñalosa, P. 2539 (12)
Peltier, J. 5692 (27)
Peña, A. 1 (24)
Pena, N. 743 (25); 764-a (36)
Pennington, T. 46-SD (30)
Pereira, E. 352 (42)
Pereira, J. 3, 35, 86, 150, 173, 224, 242, 355 (25); 588 (39); 590 (25)
Pérez, A. 1064 (30); 1473 (44); 2697 (50); 2906 (18)
Perkins, J. 1041 (49); 1042 (20); 1162 (18)
Pessoni, L. 348 (21)
Petean, M. 190, 191 (25)
Peter, A. 668, 668, 9041 (27)
Peyton, B. 439, 1285 (12)
Philipson, W. 1033, 1041, 1159, 1160 (49)
Phillips, B. 79 (49)
Pichi Sermolli, R. 4454 (27)
Picotte, C. 1765-66 (22); 1766-66, 1944-67 (9); 2083-68 (24); 2089-68 (9); 2112-68 (30)
Pignal, M. 3633 (25)
Pipoly, J. 5922 (22); 11009 (18); 11057, 14013, 15879, 16023, 16064 (44); 17770 (24)
Pirani, J. 168 (42)
Pitman, N. 256 (44); 1120 (18)
Pittier, H. 5634 (21); 7383 (4); 7568 (24); 12359 (2); 12360 (18)
Plowman, T. 4017 (44)
Plumier, C. s.n. [Herb. Tournefortianum 53] (18)
Poeppig, E. s.n. [MO-35] (21); s.n. [W-0055] (28)
Poiteau, P. s.n. [P-01409072] (18)
Polhill, R. 238, 239 (27)
Pollard, C. 114 (18); 152 (7)
Porter, D. 4670 (1); 4791 (10); 4793 (30); 4828 (10); 4937, 4938 (30); 4940 (18)
Portugal, A. 218, 235 (28); 436 (39); 446 (28); 449, 570 (12); 647 (28)
Potier, J. 187 (27)
Prado, J. 320, 530 (25); 585 (44); 1072 (25); 1117 (42); 1314 (44); 1660 (46); 2008 (25)
Prance, G. 16153 (41); 10113 p.p. (44)
Price, N. s.n. [NY-01402804] (27)
Pringle, C. 5530, 6546, 11284, 15713 (37)
Pritchard, R. 12 (30)
Pritchard, R. 43 (30); 48 (18)
Proctor, G. 3720 (49); 3979 (18); 4125, 4799, 5457 (7); 5512 (23); 5841 (20); 7637 (23); 17323 (7); 17696 (18); 17759 (7); 19104 (18);
19197, 19377 (7); 19528 (18); 20413 (24); 21700 (7); 21703 (18); 22689 (23); 25073 (34); 25660, 39836, 39894 (7); 40368 (24); 47121
(4)
Purpus, C. 250 (31); 251, 1973, 16619 (24); 1594 p.p. (37)
Purves, J. 45 (27)
• Phytotaxa 535 (1) © 2022 Magnolia Press
MATOS ET AL.
Queiroz, L. 2999 (25)
Quelal, C. 310 (44)
Quesada, F. 152 (30); 1001 (18); 1087, 1125 (2); 1418 (9); (48); 1464 (17); 1464 (24)
Questel, A. 1785, 2901 (24); 3924 (18)
Quipuscoa, V. 1037 (12); 2157 (3); 2309 (21)
Quizhpe, W. 2229 (21)
Raddi, G. s.n. [BR-000000, FI, PI-00, PI-②] (46)
Rakotondrainibe, F. 2956, 5012, 5832, 6057, 6421 (27)
Ramsden, J. s.n. [K-000804509] (27)
Ranker, T. 1544 (27)
Rankin, D. 48 (27)
Rasolohery, A. 854 (27)
Rauh, W. P1604 (44)
Raven, P. 21966 (32)
Rawson, R. 1049 (27); 3023, 3333 (18)
Regnell, A. 1302 (46); III-1438 (27)
Reitz, R. 107 (27); 266, 284, 331, 978, 1950, 2069, 3087, 4445, 6629, 12087 (25)
Reyes, J. 1174 (37); s.n. [BSC-MGC2313] (20); s.n. [BSC-MGC3357] (7)
Rezende, S. 3875 (25)
Rhoads, S. s.n. [US-03339862] (24)
Riba, R. 46 (24); 229 (18); 334 (34); 1101, 1373 (24)
Ribas, O. 716, 1348, 2169 (25)
Ridoutt, C. 11562 (3)
Riedel, L. s.n. [P-01380591] (46)
Rimachi, M. 11229 (44)
Rivera, M. 1798 (24)
Rivero, R. 1152 (4); 1616 (24); 2224 (21)
Robinson, H. s.n. [US-01565288] (18)
Robles, R. 2701 (10)
Rodas, R. 484 (24)
Rodrigues Jr., C. 1300 (46)
Rodriguez, E. 896 (44)
Rodriguéz, G. 376 (18); 332 (21)
Rodríguez, L. 1590 (44)
Rodríguez, W. 3457 (24); 4234 (30); 4250 (18); 4818 (22); 4839 (12); 5098, 5582 (22); 5587 (19); 5599 (22); 5618 (19); 5628 (22); 7058
(44)
Rohr, J. 70 (27); 329, 351 (25); 890 (27); 1012, 1017, 1030, 1066 (25)
Rojas, A. 430 (9); 513 (21); 515 (32); 611 (24); 648 (9); 1051 (22); 1094 (9); 1101 (24); 1311 (30); 1363 (9); 1440 (40); 1527, 1538, 1582
(9); 1585 (22); 1675 (40); 1691 (18); 1721 (9); 1733 (22); 1737, 1774 (24); 1786 (30); 1865 (6); 1924 (30); 1984 (40); 1991 (22); 2044
(21); 2191, 2211 (22); 2279 (18); 2280 (21); 2314 (9); 2328 (22); 2349 (6); 2360, 2390 (9); 2452 (30); 2491 (9); 2571 (30); 2591, 2605
(24); 2613 (21); 2710, 0 (17); 2812 (18); 2856 (12); 2861 (17); 2902 (6); 2907 (32); 2918 (24); 2941 (22); 2981 (18); 2983 (24); 3020
(49); 3048 (9); 3134 (22); 3141 (24); 30 (11); 3286 (9); 3311 (48); 3311 (24); 3407 (9); 3425, 3522 (24); 3573 (9); 3608 (18); 3609
(21); 30 (2); 3902, 3964, 4033 (22); 4048 (30); 4066 (24); 4084 (18); 4086 (9); 4126 (24); 4175 (1); 4182 (30); 4205 (21); 4501 (24);
4503 (9); 4525 (11); 4629 (18); 4636 (30); 4645 (24); 4786 (30); 4844 (24); 4845 (21); 4854 (22); 4919 (31); 4924 (24); 4937 (6); 4952
(9); 503 (45); 5057 (10); 5094 (17); 5102 (22); 5423 (34); 5455 (30); 5462 (18); 5472 (9); 5933 (31); 6045 (22); 30 (40); 7109, 8053
(9); 8304, 8466 (18)
Rojas, R. 108-a, 110 (44); 196 (21)
Rolleri, C. 318, 322 (27)
Rombouts, H. 871 (44)
Rose, J. 20370 (25)
Ross, H. 707 (24)
Rossi, L. 1220 (25)
Rouhan, G. 103, 107, 111, 117, 195, 214, 217, 222, 228, 250, 448 (27)
Rowlee, W. 955 (18)
A NOMENCLATOR FOR ELAPhoGLoSSUM SECTION PoLYTrIChIA Phytotaxa 535 (1) © 2022 Magnolia Press • 3
Royer, C. 72 (15); 75 (25)
Rubio, D. 1466 (30)
Rueda, R. 9182, 9591 (10)
Rusby, H. 292 (27)
Ryan, J. s.n. [BM-000335] (7)
Rzedowski, J. 23682, 25175, 26011, 44912, 46176 (37)
Saiki, Y. CR34 (30)
Salas, N. 30 (49); 55 (37); 111, 164, 171 (49)
Saldias, M. 4512 (36)
Salimena, F. 1290 (25)
Salino, A. 91, 692 (27); 1146, 1886, 1978, 2297 (25); 2502 (46); 2798 (27); 2956 (46); 4884 (36); 4896 (25); 4937 (27); 5312 (21); 5549
(36); 5852 (25); 6105 (46); 6142, 6297 (36); 6361 (21); 6442 (25); 6498 (21); 6502, 6632, 6757 (46); 7096 (27); 7328 (46); 7410 (39);
7436 (21); 7507 (25); 7707 (46); 7851 (36); 7940, 8076 (27); 8503, 8664 (25); 8834 (46); 9240 (21); 9265, 9470, 9500, 9797 (25); 9812
(39); 9934 (21); 10148 (25); 10180 (39); 10507, 10976 (27); 11048 (36); 11124 (39); 11316 (27); 11492 (25); 12345, 12562 (39); 13791
(25); 13844 (39); 14073 (15); 14137 (25); 14169 (39); 14978 (5); 15862 (31); 15846 (24)
Sánchez, C. s.n. [HAJB-HFC61481], s.n. [HAJB-HFC71330] (7); s.n. [HAJB-HFC73522], s.n. [HAJB-HFC73717], s.n. [HAJBHFC73718], s.n. [HAJB-HFC74129] (20); s.n. [HAJB-HFC75044], s.n. [HAJB-HFC75075] (7); s.n. [HAJB-HFC75124] (20); s.n.
[HAJB-HFC75172], s.n. [HAJB-HFC75173] (7); s.n. [HAJB-HFC75182] (43); s.n. [HAJB-HFC75188], s.n. [HAJB-HFC76995], s.n.
[HAJB-HFC77084], s.n. [HAJB-HFC77790], s.n. [HAJB-HFC78438], s.n. [HAJB-HFC78854], s.n. [HAJB-HFC79407] (7); s.n. [HAJBHFC80482], s.n. [HAJB-HFC80573], s.n. [HAJB-HFC81326], s.n. [HAJB-HFC81376] (20); s.n. [HAJB-HFC81396] (7); s.n. [HAJBHFC81397] (20); s.n. [HAJB-HFC81478] (7); s.n. [HAJB-HFC81502] (20); s.n. [HAJB-HFC82963] (7)
Sanchez, H. 199, 399 (37)
Sanders, A. 4409, 22572 (37)
Sandoval, Y. 71 (37)
Sandra, F. 682 (49)
Sanín, D. 5077 (21)
Sanoja, E. 3026 (21)
Santos, N. s.n. [R-52117] (25)
Sargent, F. A16 (18)
Scamman, E. 6006, 6007 (9); 6541 (7); 6542 (18); 7161 (9); 7167 (24); 7700 (9)
Schaffner, W. (31); 30, 281 (24)
Schelpe, E. 2549, 2603, 2624, 2798, 5523, 5530, 5580 (27)
Schipp, W. 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 •