10 11646@phytotaxa 426 1 1

Phytotaxa 426 (1): 001–105 ISSN 1179-3155 (print edition)
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Monograph PHYTOTAXA
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https://doi.org/10.11646/phytotaxa.426.1.1
PHYTOTAXA
426
Synopsis of the genus Phlegmariurus (Lycopodiaceae) in

Colombia
BENJAMIN ØLLGAARD
Herbarium AAU, Science Museums, and Section for Ecoinformatics and Biodiversity, Institute for Bioscience, Building 1137,
University of Aarhus, DK-8000 Aarhus, Denmark.
benjamin.oellgaard@bios.au.dk
Magnolia Press
Auckland, New Zealand
Accepted by Marcus Lehnert: 24 Oct. 2019; published: 16 Dec. 2019 1

BENJAMIN ØLLGAARD
Synopsis of the genus Phlegmariurus (Lycopodiaceae) in Colombia
(Phytotaxa 426)
105 pp.; 30 cm.
16 December 2019
ISBN 978-1-77670-837-6 (paperback)
ISBN 978-1-77670-838-3 (Online edition)
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ISSN 1179-3155 (print edition)

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2 • Phytotaxa 426 (1) © 2019 Magnolia Press ØLLGAARD

CONTENTS
Abstract ...........................................................................................................................................................................................................3
Introduction .....................................................................................................................................................................................................3
Terminology ....................................................................................................................................................................................................4
Group of Phlegmariurus funiformis................................................................................................................................................................5
Group of Phlegmariurus myrsinites................................................................................................................................................................6
Group of Phlegmariurus acerosus ................................................................................................................................................................15
Group of Phlegmariurus dichotomus............................................................................................................................................................20
Group of Phlegmariurus hartwegianus ........................................................................................................................................................28
Group of Phlegmariurus lindenii ..................................................................................................................................................................32
Group of Phlegmariurus taxifolius ...............................................................................................................................................................33
Group of Phlegmariurus linifolius ................................................................................................................................................................35
Group of Phlegmariurus reflexus .................................................................................................................................................................43
Group of Phlegmariurus firmus ....................................................................................................................................................................52
Group of Phlegmariurus tetragonus .............................................................................................................................................................57
Group of Phlegmariurus crassus ..................................................................................................................................................................59
Excluded names ..........................................................................................................................................................................................101
Acknowledgements .....................................................................................................................................................................................101
References ...................................................................................................................................................................................................101
Abstract
This article presents the results of a study of a large material of Colombian Lycopodiaceae, describing 75 species of the
genus Phlegmariurus, some of them subdivided into several varieties. Their nomenclature, distribution and habitats are indi-
cated, and problems of species delimitation and infraspecific variation are discussed. All species are illustrated, and studied
specimens are cited. An attempt has been made to group the species, mainly following the clades of recent molecular stud-
ies. The following taxa are described as new to science: Phlegmariurus cocuyensis, P. cruentus var. exilis, P. cruentus var.
illimitatus, P. idroboi, and P. josesantae.
Keywords: Lycopodiaceae, Phlegmariurus, Colombia
Introduction
Colombia probably has the highest species diversity of Lycopodiaceae (P.Beauv. ex Mirbel in Lam. & Mirbel) in South
America, although in relation to its area it is perhaps equalled by Ecuador. Earlier accounts based mainly on material
in Colombian herbaria (Murillo-Pulido & Murillo-Aldana 1999; Murillo-Pulido,Murillo-Aldana, & León-Parra, 2008)
included 42 species of Phlegmariurus (as Huperzia) and 62 species of Phlegmariurus (as Huperzia) respectively.
Some regional Colombian studies: Ramírez Padilla & Macias Pinto (2007) included 29 species (as Huperzia) in a
catalog of the species in Departamento Cauca. Vargas Gaviria, Buitrago & Esquivel, (2018) recorded 10 species (as
Huperzia) from the Anaime páramo in Departamento Tolima .
For comparison, a recent count of the Lycopodiaceae for Ecuador by Øllgaard (2016b) made a total of 66 species of
Phlegmariurus. The present study includes 75 species for Colombia, several of which were described as a preparation
for the present paper.
The two Colombian accounts (Murillo-Pulido & Murillo-Aldana 1999, Murillo-Pulido, Murillo-Aldana, & León-
Parra (2008) followed the generic concept of Øllgaard (1987) with three genera Lycopodium, Lycopodiella and
Huperzia. The genera Lycopodium and Lycopodiella are now commonly subdivided into several segregate genera (see
Øllgaard 2012b , Øllgaard & Windisch 2014) and PPG (2016). The genus Huperzia s. lat. following Øllgaard (1987)
is now commonly divided into Huperzia s. str. and Phlegmariurus. No species of Huperzia s. str. have been recorded
for Colombia, so all Huperzia-species of the earlier accounts are here placed in the genus Phlegmariurus. The genus
Huperzia differs from Phlegmariurus by the production of bulbils for vegetative reproduction and spores with concave
outline between truncate corners, as opposed to convex outline between the obtuse corners in Phlegmariurus.
Phylogenetic studies (e.g. Wikström, Kenrick & Chase 1999, Testo, Øllgaard, Field, Almeida, Kessler & Barrington
2018) of Huperzia s. lat. (including Phlegmariurus) indicate that while the species of Huperzia s. str. are primary
terrestrial, the terrestrial species of Phlegmariurus have been derived from epiphytic species. The diversification of
SYNOPSIS OF THE GENUS PHLEGMARIURUS Phytotaxa 426 (1) © 2019 Magnolia Press • 3
this large group of secondarily terrestrial Phlegmariurus species in Colombia is likely to have been triggered by the
Andean uplift in the Mid Miocene no more than 15 million years ago (Wikström & Kenrick 2001, Field, Testo, Bostock
& Waycott 2015 , Testo, Øllgaard, Field, Almeida, Kessler & Barrington 2018). The youth of the High Andine group of
Phlegmariurus is reflected in poor resolution of the cladograms involving the species of this group (e.g. Wikström &
Kenrick 2000, Testo, Øllgaard, Field, Almeida, Kessler & Barrington 2018), and in several cases of highly problematic
species definition, probably reflecting habitat instability during the vegetation history of the Andean uplift as described
in numerous publications by Thomas van der Hammen and collaborators (e.g. van der Hammen, 1974). They explain
the complex vegetation history during the Quaternary, with vegetation zones moving up and down the mountains
reflecting climate changes, causing mountain top plant populations to become isolated during warm periods and to
merge and mix during cold periods.
The present study involves more than a thousand Colombian collections from most major herbaria in Europe,
North America and Colombia. Extended loans from the following herbaria were generously made available for this
study, in addition to generous gift material in AAU: AWH, B, BM, BONN, BP, BR, C, COL, CUVC, E, F, FI, G, GH,
GL, HBG, HUA, K, L, LG, LOJA, M, MA, MEDEL, MER, MICH, MO, MSC, NO, NY, OXF, P, PRC, QCA, QPLS,
RB, S, SI, U, UC, UPS, US, VEN, W, and Z (acronyms following Thiers 2019 ).
Not least the important collections by José Cuatrecasas, Antoine M. Cleef, and José Santa contributed to a broad
and representative picture, and at the same time exposed the taxonomic problems in the genus in Colombia, particularly
in the high Andes.
Terminology
Branching patterns:—Øllgaard (1979) found three basic branching patterns in the Lycopodiaceae, corresponding
to three subfamilies . In subfamily Huperzioideae Huperzia and Phlegmariurus are characterized by isotomy, i.e.,
all dichotomies result in branches of equal thickness. Isotomous branch pairs may be homoblastic or heteroblastic.
Heteroblasty, i.e., the differentiation of isotomous branches to distinct functions and aspects, occurs in many terrestrial
high Andean species of Phlegmariurus. In such plants, individuals have prostrate or even subterranean, rooting shoots
from which erect, aerial shoots bearing sporangia arise. The degree of heteroblasty appears to be affected by the
character of the substrate in the habitat. Heteroblasty is usually most pronounced in plants growing on solid soil, while
in deep, soft, e.g. mossy substrate, the prostrate shoots tend to become ascending and with fewer or no roots. Many
herbarium collections do not include the prostrate divisions in species where they are expected, making identification
troublesome sometimes. Further, with age, the prostrate, rooting shoots may gradually become ascending to nearly
erect and no longer produce roots. This evidently affects the significance of heteroblasty as a key character. Homoblasty
is the condition when isotomous branches delvelop equal functions and aspects.
Subfamilies Lycopodioideae (In Colombia: Austrolycopodium, Diphasiastrum, Diphasium, Lycopodium) and
Lycopodielloideae (in Colombia: Lycopodiella, Palhinhaea and Pseudolycopodiella) have anisotomous branching,
i.e., dichotomies result in unequally thick branches (=pseudomonopodial growth, according to Wagner & Beitel, 1992).
In Lycopodioideae, the ramifications are dorsolateral, whereas in Lycopodielloideae they are flabellate in horizontal
shoots, and dorsal in erect, aerial, strobiliferous shoots. The erect aerial shoots in Palhinhaea are dorsal in origin and
repeatedly ramified forming several flabellate lateral branchlet systems that may ultimately be strobiliferous.
Heterophylly and homophylly:—–Heterophylly and homophylly were often used as the key basis for the
classification of the species belonging to Huperzia and Phlegmariurus in several earlier treatments (Spring 1842,
1849; Baker 1887; Pritzel 1901; W. Herter 1909a; G. Herter 1949, 1950). In these classifications, the species were
keyed as either completely homophyllous (i.e., all leaves uniform), gradually heterophyllous (i.e., with gradually
reduced leaves toward the apex of the plant), or heterophyllous (i.e., with sharply distinct, long, expanded leaves in the
basal divisions, and small, reduced leaves in the narrow terminal divisions). These characters are variable, even within
a single species, and are therefore unsuitable to define higher levels of classification. For instance, several of the clades
followed in the groups below include both homophyllous and heterophyllous species. However, they are still useful for
species identification, when multiple key entries are provided.
In the following the Colombian species are arranged mainly according to major clades presented by Testo et al.
(2018). Species not included in that study are placed following the judgement of the present author.
The species treatments include nomenclature and synonymy, reference to published illustrations, and/or to original
illustrations in the present article, a description, comments on distribution, habitats, notes on variation, related species
or taxonomic problems, and citation of representative specimens.
Group of Phlegmariurus funiformis
Phlegmariurus funiformis (Spring) B.Øllgaard (2012b: 15). Lycopodium funiforme Spring (1841: 516). Huperzia
funiformis (Spring) Trevisan (1874: 248). Urostachys funiformis (Spring) Herter (1925: 387). Type: GUADALUPE:
L’Herminier s. n. (lectotype BR, GH, NY, P, RB, UC, designated by Spring (1849: 22).—Fig. 1D
Illustrations:—Øllgaard (1988: Figure 17D); Lellinger (1989: Figure 25).
FIGURE 1. A. Phlegmariurus phylicifolius, (Camp 4235A, Ecuador, AAU) growth habit. A1. Close-up of basal division. A2. Close-up
of sporangiate division.—B. Phlegmariurus subulatus (Camp 5189, Ecuador, AAU), growth habit. B1. Close-up of basal division, B2.
Close-up of sporangiate division. C. Phlegmariurus lindenii (Ecuador: Holm-Nielsen et al. 5900, AAU) C1. Close-up of basal division.
C2. Close-up of sporangiate division. D. Phlegmariurus funiformis (Ecuador: Asplund 19290, S). D1. Close-up of sporangiate division.
Del. B. Johnsen
Plants robust, flaccidly pendulous, rope-like, at least to 250 cm long. Shoots homophyllous, almost equally thick
throughout, 5–10(–15) mm in diameter including leaves, or sometimes tapering from a thicker base with patent-
ascending leaves. Stems excluding leaves 1.5–3(–5) mm thick at the base, ridged by decurrent leaf bases, pale greenish
to brownish, usually sporangiate from 40–100 cm above the base and upward, at least to 5 times dichotomous. Leaves
almost uniform throughout, or slightly shorter upward, borne in alternating whorls of 7–8, these 1.5–3 mm apart, forming
14–16 often indistinct longitudinal ranks, densely covering the stem, usually closely falcate-appressed throughout, not
twisted, linear-subulate, widest just above the base, 6–10(–12) × 1–1.5 mm, evenly tapering into a long pungent apex,
firmly herbaceous to coriaceous, dull to shining, abaxially strongly convex, often apically conduplicate, with smooth
margins. Sporangia 1–1.5 mm wide.
Distribution:—West Indies, southern Mexico to Panama, Guyana, Venezuela to Peru.
Habitat:—Epiphytic in lower montane rain forest, and gallery forest, elev. 0–1400 m.
Notes:—The name of Lycopodium funiforme was first given by Bory de Saint-Vincent, in Brongniart, Hist. Veg.
Foss. 2: 10 (1837) as a nomen nudum, and Bory was later erroneously given as the author of the name in several
publications. Spring (1849: 22) excluded a dubious Chamisso collection, labelled “California” (Herb. Bory, P), and
thus indirectly selected the l’Herminier specimen as the lectotype.
Phlegmariurus funiformis is a remarkable species, being a large, pendulous epiphyte with a distinctive, ropelike
aspect, due to the very regularly appressed, coriaceous, strongly convex, subulate leaves. not easily confused with
other species. Molecular studies (Wikström & Kenrick 2001: 157), Field et al. (2015: 11) and Testo et al. (2018)
indicate that this species belongs to a paleotropical clade without relatives in the Neotropics.
Specimens examined:—Antióquia: Mun. Urrao, Vereda Calles, Bosque Nacional Natural Las Oequideas;
Quebrada Honda; ridge NW of Cabaña Calles, 1330 m, Pipoly et al. 16920 (AAU). Cauca: Chuaré, 100 m, 23
Dec 1946, Haught 5374 (US). Nariño: Junin, 850 m, Foster 2180 (GH). Quebrada Cartagena, near Ricaurte, Río
Guabo drainage, 1220 m, Ewan 16887 (NO, US). Valle del Cauca: Río Calima (reg. del Chocó), La Trojita, 5–50 m,
Cuatrecasas 16553 (COL, F, GH, US). Buenaventura, corregimiento de San Isidro, 30–40 m, Santa & Sánchez 1048
(AAU, COL, HUA).
Group of Phlegmariurus myrsinites
This group includes mainly rather delicate pendulous and heterophyllous epiphytic species, with the surprising
exception of the homophyllous Phlegmariurus capillaris, earlier assumed to be a close relative of P. linifolius.
Phlegmariurus amentaceus (B.Øllg.) B.Øllgaard (2012b: 53). Lycopodium amentaceum B.Øllg. (1982: 53).—Huperzia
amentacea (B.Øllg.) Holub (1985: 70). Type:—VENEZUELA: Est. Mérida: En la faja superior de la selva montanera,
3000 m, Camino al Páramo Los Monsalves, Vareschi & Pannier 1817 (holotype VEN, isotypes M, VEN).—Fig. 2
Illustrations: Øllgaard (1982: 53).
Delicate, pendulous, to 60 cm long. Shoots heterophyllous. Basal divisions with expanded leaves, 10–20 mm in
diameter including leaves, terminal divisions abruptly constricted to 1.2–1.8 mm thick. Stems excluding leaves 0.6–1.3
mm thick at the base, greenish to bright red. Expanded leaves decussate or subdecussate, spreading to perpendicular,
straight to slightly recurved, oblong to oblong-lanceolate, usually widest at the middle, obtuse or mucronulate, (4–)6–
10 × 2–3(–3.5) mm, flat, entire. Leaves of terminal constricted divisions decussate or subdecussate, appressed and
clasping with their bases, widely ovate to subcordate, 1.2–1.7 × 1.2–1.5 mm, abaxially rounded to carinate, equalling
or slightly exceeding the sporangia. Sporangia ca. 1 mm wide.
Distribution:—Guatemala, Venezuela, Colombia.
Habitat:—Epihytic in moist forest, elev. 1800–3000 m.
Notes:—Related to Phlegmariurus callitrichifolius and P. cuneifolius. These species are mainly separated on the
basis of subtle differences in the shape and size of the expanded leaves.
Spcimens examined:—Cundinamarca: Cerro Negro, Hato Grande, ridge E of Río Muchindote, 15 km E of
Gachetá, Cordillera Oriental, 3000 m, Grant 9436 (COL, US); Road to the E from Guasca, 3200 m, Haught 5826
(COL, US). Monserrate, Bogotá, 2900 m, Lindig 1504 (K, P).

FIGURE 2. Phlegmariurus amentaceus (Colombia: Haught 5826, COL) A. Growth habit. A1. Details of terminal, including sporangiate,
divisions.
Phlegmariurus aqualupianus (Spring) B.Øllgaard (2012a: 480). Lycopodium aqualupianum Spring (1841: 518).
Urostachys aqualupianus (Spring) Herter (1923: 166).—Huperzia aqualupiana (Spring) Rothmaler (1944: 62).
Lectotype:—GUADELUPE: L’herminier (lectotype LG, designated by Badré 1983: 4; isolectotypes BM, K, P).—
Fig.3.
Plants pendulous, flaccid, 3–4 times dichotomous. Stems stramineous, angular, 15–50 cm long, thin, usually less than
1 mm thick excluding leaves. Shoots dimorphic, the proximal divisions with wide expanded leaves, the distal divisions
abruptly narrowed, with small imbricate leaves. Proximal expanded divisions with uniform leaves, 10–17 mm wide
including the leaves, sometimes slightly tapering distally, usually continuously overlapping (pressed). Expanded
leaves usually decussate (sometimes in alternating whorls of 3), forming 4(–6) longitudinal rows, regularly inserted,
ascending to patent, softy herbaceous, ovate to elliptic-oblong, 6–11 × 2.8–4 mm, slightly apiculate, with decurrent
base. Constricted divisions quadrangular, with extensive sterile zones proximally. Leaves of constricted divisions
imbricate, regularly decussate, ovate, carinate and usually long-acuminate, 3–4.5 ×1.5 mm. Sporangia ca. 1 mm wide,
completely concealed by sporophyll bases.
Distribution:—West Indies, Colombia, Venezuela, Guyana, Brazil.
Habitat:—Epiphytic in wet premontane and cloud forest, 1200–2400 m.
Notes:—This species is related to Phlegmariurus ericifolius (C. Presl) B.Øllgaard (Ecuador to Bolivia) and P.
dichaeoides. P. ericifolius can usually be distinguished by its narrower, more acute expanded leaves in whorls of 3,
while P. aqualupianus usually has ovate, subacute to obtuse, usually decussate expanded leaves.
Specimens examined:—Antioquia: Mun. Guatapé, near San Rafael, 1890 m, de Escobar & Folsom 2409 (HUA).
Road Medellín–Bogotá, Santuario, 2000–2100 m, Uribe 6764 (COL). La Quebra, road to Yolombó, 2000 m, Lehmann
XLIVa (G). Cundinamarca: Sasaima, San Bernardo at Río Guane, 1500 m, Schneider 610 (S). Magdalena: Sierra
Nevada de Santa Marta, Alto Río Buritaca, 1690 m, Jaramillo et al. 5139A (COL). Meta: Cord. de Macarena, extreme
NE, Renjifo, 1300–1900 m, Idrobo & Schultes 994 (AAU, COL, GH, US); Sierra de la Macarena, central mountains,
N ridge, 1500 m, Philipson & Idrobo 2012 (COL, US). Santander: Cerro Armas, 1200–1400 m, Haught 1954 (COL,
GH, US); Mesa de Los Santos, 1500 m, Killip & Smith 15287 (US). Putumayo: Las Mesas, near Pepino, 1370 m,
Foster 2234 (COL, GH). Valle: Cord. Occ., W slope, Hoya del Sanquinini, La Laguna, 1250–2000 m, Cuatrecasas
15611 (F, US). Finca Kyburz, E slope above Río Bitaco, 1 km E of Bitaco, Pacific slope of the W Cord., 1350–1400
m, Hutchison & Idrobo 3063 (F, GH, M, MO, NY, P, UC, US). La Cumbre, 2100–2400 m, Killip 11383 (GH, NY, US ).
FIGURE 3. Phlegmariurus aqualupianus (Colombia: Idrobo 994, AAU). A. Growth habit. A1. Details of terminal, including sporangiate,
divisions.

Phlegmariurus callitrichifolius (Mettenius) B.Øllgaard (2012b: 13).Lycopodium callitrichifolium Mettenius (1865:
309)—Urostachys callitrichaefolius (Mettenius) Nessel (1939: 253)—Huperzia callitrichifolia (Mettenius) Holub
(1985: 71). Type:—COLOMBIA: Bogotá, Tabano, Triana s.n. [ex herb. Mettenii] (holotype B, isotype P).—Figure
4C.
Illustrations:—Øllgaard (1988: Fig. 23B).
Delicate plants, pendulous, at least to 60 cm long. Shoots heterophyllous, in the basal divisions, to 30 cm from the base,
(5–)7–15 mm in diameter including the expanded leaves, then abruptly constricted to 1–2 mm in diameter including
the reduced, imbricate leaves in the terminal, constricted, quadrangular divisions. Stems excluding leaves 1 mm thick,
or less, at the base, tapering to ca. 0.5 mm, greenish to bright red, at least to 10 times dichotomous. Expanded leaves
of basal divisions decussate or subdecussate, the leaf pairs 2–4 mm apart, forming 4 longitudinal ranks, spreading
to perpendicular or slightly reflexed, oblong to widely ovate, widest at or below the middle, obtuse or mucronulate,
3.5–6(–8) × 2.5–3.5 mm, flat, the lamina twisted to a vertical position, softly to firmly herbaceous. Leaves of terminal
constricted divisions decussate, often sporangiate in rather short zones of the divisions, appressed and clasping with
their bases, widely ovate to subcordate, 1.5–2 × 1–1.5 mm, carinate, with falcate to reflexed mucronulate tips, the
sporophylls equalling or slightly exceeding the sporangia. Sporangia ca. 1 mm wide.
Distribution:—Colombia to Northern Peru.
Habitat:—Cloud forest, epiphytic or hanging from mossy banks, elev. 2500–3200 m.
Notes:—Phlegmariurus callitrichifolius is closely related to P. cuneifolius and P. amentaceus.
Ramírez Padilla & Macias Pinto (2007) reported this species from Dept. Cauca but I have not been able to encounter
any specimen from this Departamento.
Specimens examined:—Province unknown: Tabano, Andre K 1464 (K). Cundinamarca: Salto del Tequendama,
2500 m, Bischler, H. 2267 (COL). Tolima: Road Mariquita–Manizales, La Libia, 3000–3200 m, Santa, J. & L. de
Escobar 853 (AAU, COL, HUA).
FIGURE 4. A. Phlegmariurus cuneifolius (Peru, Hutchison & Wright 6943, P) growth habit. A1. Close-up of basal division. A2. Close-up
of sporangiate division.—B. Phlegmariurus dichaeoides (Ecuador: Cazalet & Pennington 5114, US) growth habit. B1. Close-up of basal
division, B2. Close-up of sporangiate division.—C. Phlegmariurus callitrichifolius (Ecuador: Holm-Nielsen et al. 3988, AAU) growth
habit. C1. Close-up of basal division. C2. Close-up of sporangiate division. Del B. Johnsen.
Phlegmariurus capillaris (Sodiro) B.Øllgaard (2012a: 480).Lycopodium capillare Sodiro (1883: 90). Lycopodium
sarmentosum Spring var. capillare (Sodiro) Sodiro (1893: 572). Urostachys capillaris (Sodiro) Herter (1949: 54).
Huperzia capillaris (Sodiro) Holub (1985: 71). Type:—ECUADOR (Prov. Pichincha ?):, Hda. Guanaxilla, 480 m,
Sodiro s. n. (holotype K). The type locality is probably in the area NE of Santo Domingo de los Colorados.—Fig. 5C.
Lycopodium sarmentosum Spring var. rubescens Spring (1849: 13). Urostachys rubescens (Spring) Herter (1949: 79). Type:—COLOMBIA:
prov. Popoyan, Hartweg 1464 (K with Spring’s annotation: lectotype B; isotypes P, NY fragment).
Lycopodium underwoodianum Maxon (1909: 41).Urostachys underwoodianus (Maxon) Herter (1949: 87). Huperzia underwoodiana
(Maxon) Holub (1985: 77). Type:—COSTA RICA: vicinity of Coliblanco, about 1950 m, Maxon 213 (holotype US, isotypes BM,
C, GH, NY, P).
Lycopodium guatemalense Maxon (1913: 177). Urostachys guatemalensis (Maxon) Herter (1923: 165). Type:—GUATEMALA:
Pansamalá, Tuerckheim, ed. Donn.-Sm. 957 (holotype US; isotype UC).
Illustrations: Lellinger (1989: Fig. 17); Øllgaard (1988: Fig. 19C).
FIGURE 5. A. Phlegmariurus polycarpos (Ecuador, Sodiro s.n., UC) growth habit. A1. Close-up of sporangiate division.—B.
Phlegmariurus dichotomus, (Holdridge 1654, Ecuador, GH) growth habit. B1. Close-up of sporangiate division.—C. Phlegmariurus
capillaris, (Hartweg 1464, Colombia, P) growth habit. C1. Close-up of sporangiate division. Del. B. Johnsen

Very delicate, pendulous, with flaccidly hanging divisions, 9–50 cm long. Shoots homophyllous, rarely gradually
heterophyllous, 7–15 mm in diameter including leaves in basal divisions. Stems excluding leaves 0.3–0.5(–0.8) mm thick
near the base, slightly tapering upward, straight or somewhat flexuous, usually bright red, discontinuously sporangiate
from 6–25 cm above the base and upward, to 6(–13) times dichotomous. Leaves almost uniform throughout, borne
apparently randomly, spirally arranged, occasionally in pairs or whorls of 3, 1–3(–5) mm apart, spreading, straight
to falcately curved, with strongly twisted, often deflexed lamina base, linear-lanceolate, widest in the middle or the
basal half, 5–10(–12) × 0.4–0.7(–1) mm, almost flat, or slightly convex adaxially, soft-herbaceous, with obscure to
somewhat prominent vein above. Lamina base tinged with red. Sporangia 0.7–1 mm wide.
Distribution:—Central America, Venezuela, Colombia, Ecuador, Peru, Bolivia Brazil.
Habitat:—Epiphytic, in lower montane rain forest, 850–2440 m.
Notes:—Earlier assumed to be closely related to Phlegmariurus linifolius. Differs in the more delicate growth
habit. Stem coloration is variable, the red colour lacking in some collections. Rather surprisingly, its molecular data
point to a relationship to P. phylicifolius (Testo et al. 2018).
Specimens examined:—Antioquia: Antioquia 1943, Kalbreyer s.n. (K, NY fragm.). Cauca: La Gallera,
Micay Valley, Cord. Occid. 1400–1500 m, Killip & Smith 7741 (GH, US). Popoyan, Hartweg 1464 (B, K, P). Huila:
Mountains 6 km S of Cueva de los Guácheros, ca. 20 km SW of Alejandría, 2440 m, Little 8506 (COL, GH, US). Norte
de Santander: Ocaña, 2440–3050 m, Schlim 507 (BR, G, K).
Phlegmariurus cuneifolius (Hieron.) B.Øllgaard (2012b: 14). Lycopodium cuneifolium Hieronymus (1905: 572).—
Urostachys cuneifolius (Hieron.) Nessel (1939: 252).—Huperzia cuneifolia (Hieron.) Holub (1985: 72). Type:—
COSTA RICA. Heredia: Volcán Barba, Hoffman 50 (lectotype B, designated by Nessel (1939: 252); isotype NY
[fragment]).—Fig. 4A.
Pendulous, very delicate, to 60 cm long. Shoots heterophyllous. Basal divisions (5–)7–13 mm in diameter including
the expanded leaves, abruptly constricted to 1–2 mm in diameter in terminal, constricted, quadrangular divisions
with imbricate leaves. Stems excluding leaves 1 mm thick or less at the base, greenish to bright red. Expanded leaves
of basal divisions decussate, the leaf pairs 2–4 mm apart, spreading to perpendicular, oblanceolate to obovate or
spathulate, cuneate, widest at or above the middle, obtuse or mucronulate, 3–6 × 1.5–3 mm. Leaves of terminal
constricted divisions often sporangiate in rather short zones of the divisions, decussate, appressed and clasping, widely
ovate to subcordate, 1–1.5(–2) × 1–1.5 mm, carinate, the sporophylls equalling or slightly exceeding the sporangia.
Sporangia ca. 1 mm wide.
Distribution:—Costa Rica, Panamá, Andean Venezuela, Colombia, northern Peru.
Habitat:—Cloud forest, epiphytic or hanging from mossy banks, 2300–3400 m.
Notes:—Closely related to Phlegmariurus callitrichifolius and P. amentaceus.
Specimens examined:—Antioquia: Kalbreyer 514 (K). Kalbreyer 773 (K). Arauca: San José Valley, below El
Playón, near the old trail to Llanos (San Lope), 3400 m, Grubb & Guymer P 111 (US). Boyacá: Sierra Nevada del
Cocuy, San José Valley, below El Playon, near trail San Lope, Grubb & Guymer (P. 111 (photo ex US). Guajira: Sierra
Nevada, Río Hacha, 2440 m, Schlim 881 (BR, P). Norte de Santander: Ocaña, 2440–3050 m, Schlim 468 (BR, P).
Santander: E Cord., S slope of Mt. San Martín, near Charta, 2300–2500 m, Killip & Smith 19198 (GH, NY, US).
Illustrations: Lellinger (1989: Fig. 21); Øllgaard (1988: Fig. 23A).
Phlegmariurus dichaeoides (Maxon) B.Øllgaard (2012b: 14). Lycopodium dichaeoides Maxon (1905: 231).—
Urostachys dichaeoides (Maxon) Nessel (1939: 252).—Huperzia dichaeoides (Maxon) Holub (1985: 72). Type:—
GUATEMALA: Alta Verapaz, near Finca Sepacuite, Cook & Griggs 251 (holotype US).—Fig. 4B.
Illustrations: Lellinger (1989: Fig. 22); Øllgaard (1988: Fig. 23C).
Plants slender, pendulous, at least to 60 cm long. Shoots heterophyllous, in the basal divisions, to 25–35 cm from the
base, 12–17 mm in diameter including the wide, expanded leaves, then abruptly constricted to 1–2 mm in diameter
including the imbricate, reduced leaves of the 10(–16) cm long sporangiate, terminal divisions. Stems excluding
leaves ca. 1 mm thick, tapering to ca. 0.5 mm, pale greenish, to 7 times dichotomous. Expanded leaves uniform
in shape, size, and position throughout, in 4 regular ranks, decussate, nearly continuously overlapping throughout
(pressed specimens), diverging 65°–90° from the stem, the leaf pairs 2–3 mm apart, leaves elliptic to oblong (rarely
suborbicular), widest in the middle or just below, 6–9 × 3–4 mm, with mucronulate apex and flat to slightly revolute
margins, the vein obscure above, evident but not prominent below, the lamina usually twisted to a vertical position.
Leaves of apical constricted divisions decussate, all, or almost all, sporangiate, appressed and clasping with their bases,
ovate to triangular-ovate, short- to long-cuspidate, somewhat involute, sharply carinate, at least in the apex, 1.2–1.5(–
2.7) × 1(–1.3) mm, equalling the sporangia, or to 1.5 times longer, the distance between leaf pairs 0.5–1(–2.5) mm.
Sporangia 0.7–1 mm diam.
Distribution:—Guatemala to Panama, Pacific slopes of the Andes of Colombia and Ecuador. Ramírez Padilla &
Macias Pinto (2007) reported this species from Dept. Cauca.
Habitat:—Epiphytic in lower, wet montane forest, 100–1700 m.
Notes:—Differs from Phlegmariurus aqualupianus by the widely elliptic, wide-spreading to nearly perpendicular,
less continuously overlapping, and the short, less sharply angular constricted divisions with short-tipped imbricate
leaves. Seems to replace Phlegmariurus aqualupianus in the western part of Colombia.
Specimens examined:—Province unknown: Juntas, Lehmann BT 1164 (NY, US). Antioquia: Mun. Guatapé/
San Rafael, 1890 m, Santa & Vallejo 1070 (AAU). Urrao: P.N. Las Orquídeas, Vereda Calles, 1450–1500 m, Pipoly et
al. 17344 (MO). Urrao: P.N. Las Orquídeas, Vereda Calles, 1450 m, Pipoly et al. 17172 (MO). Chocó: Límit between
Colombia y Panamá, serranía del Darién, 1400 m, Gentry et al. 16850 (MO). 0.5 kms N de San José del Palmar,
Lellinger & de la Sota 733 (US). Valle del Cauca: Old road Cali–Buenaventura, drainage of río Digua, Piedra de
Moler, 900–1180 m, Cuatrecasas 14953 (US). Old road Cali–Buenaventura, 9 km below Central Hidroeléctrica. del
Anchicayá, 300 m, de Escobar et al. 2621 (HUA).
Phlegmariurus ericifolius (Presl) B.Øllg.
was reported (as Huperzia) by Ramírez Padilla & Macias Pinto (2007) Dept. Cauca, without specimen citation. The
presence in Colombia is likely, as the species is known from northernmost Ecuador in Prov. Carchi.
Illustration: Øllgaard (1988: Fig. 23D).
Phlegmariurus myrsinites (Lamarck) B.Øllgaard (2012b: 17).—Fig. 6.
Lycopodium myrsinites Lamarck (1792: 654).—Plananthus myrsinites (Lamarck) P. Beauvois (1805: 111).—Urostachys myrsinites
(Lamarck) Herter (1923: 166).—Huperzia myrsinites (Lamarck) Trevisan (1874: 249). Type:—HAITI: “S. Domingue”, Comm.
Joseph Martin (holotype P [Herb, Lamarck ], isotype P-00559116).
Lycopodium patens Willdenow ex Sprengel (1827: 12).—Urostachys patens (Sprengel) Nessel (1939: 2345). Type:—Provenance
unknown: anonymous (holotype B-Willdenow 19342).
Lycopodium roraimense Underwood & Lloyd (1906: 115).—Urostachys roraimensis (Underwood & Lloyd) Nessel (1939: 246). Type:—
BRITISH GUIANA: forest slopes near Roraima, Herb. Jenman (holotype NY).
Lycopodium skutchii Maxon (1933: 159). Type:—GUATEMALA: Chimaltenango, Chichavac, elev. 2400–2700 m, Skutch 243 (holotype
US-1494904).
Illustrations: A.R. Smith (1981: Fig. 82 a, b); Lellinger (1989: Fig. 32); Mickel & Beitel (1988: Fig. 3A).
Description:—Plants pendulous, at least to 65 cm long. Shoots heterophyllous, usually not all sharply differentiated in
the basal divisions 10–18 mm in diameter including the expanded leaves, upward gradually, within a short transition, or
abruptly constricted to 1.5–3 mm thick including the reduced, imbricate leaves in the terminal divisions. Stem excluding
leaves ca. 1 mm thick at the base, tapering to ca. 0.5 mm, pale greenish, to 7 times dichotomous. Expanded leaves
of basal divisions decussate or subdecussate, often irregularly shaped, subdistant to densely crowded and somewhat
overlapping, the leaf pairs 1.5–6 mm apart, ascending to perpendicular, ovate-lanceolate or narrowly so, acute, usually
the widest ones with a rounded base, 6–11 × 1.5–3 mm, usually flat, straight to somewhat recurved, the lamina twisted
to a vertical position. Leaves of terminal constricted divisions highly variable, often with complete reduction series,
and with recurrent series to expanded shape, decussate or subdecussate, continuously or discontinuously sporangiate.
Transitional leaves with widely ovate base, and short to long acuminate apex, appressed and clasping, with the wide
base abaxially rounded to bluntly carinate, with straight to recurved apex, 2.5–5 × 1.5–2 mm. Shortest leaves with
base conform, but with straight to falcate apex, bluntly to sharply carinate, scarcely exceeding the sporangia, ca. 2 mm
long. Sporangia 1–1.3 mm wide.
Distribution:—Hispaniola, S Mexico to Costa Rica, Trinidad, Hispaniola, British Guiana, Venezuela to Bolivia.
Habitat:—Epiphytic in moist forest, lower Pacific slopes of the Andes, 1300–ca. 2500 m

FIGURE 6. Phlegmariurus myrsinites (Venezuela: Meier & Silva 1585, AAU). A. Growth habit. A1. Detail of sporangiate divisions.
Notes:—Phlegmariurus myrsinites is related to P. phylicifolius. Forms of the latter, described as Lycopodium

subulatum var. pastoense Baker resemble P. myrsinites in the gradual transition, and often repeated reversion from
expanded to imbricate, reduced leaves. In Colombia the two species may be separated mainly by the wider, softer,
and thinner expanded leaves, and larger sporangiate leaves of P. myrsinites. Species of the group of Phlegmariurus
quadrifariatus (Bory 1829: 245) B.Øllgaard (2012a: 481) (SE Brazil) were erroneously placed under P. myrsinites by
several earlier authors (e. g. Nessel 1927, 1939).
Specimens examined:—Cauca: 3 km E of Timbo, 2135 m, Drew E-782 (US). Magdalena: Santa Marta, Cerro
de Onaca, Smith 2469 (NY).
Phlegmariurus phylicifolius (Poiret) B.Øllgaard (2012b: 17).—Fig. 1A.
Lycopodium phylicifolium Desvaux ex Poiret (1814: 546).—Urostachys phylicifolius (Poiret) Nessel (1939: 246).—Huperzia phylicifolia
(Poiret) Holub (1985: 75). Type:—CHILE. Prov. unknown: “Habitat in Chili,” Herbier de A.N. Desvaux, donnée par Mme Vve.
Lavallée en 1896 (lectotype P00559236, designated by Øllgaard (1988: 117).
Lycopodium congestifolium Spring (1842: 70). Type:—PERU: Without locality, Dombey, H. M. P., Herb. Deless. (holotype P).
Lycopodium subulatum Poiret var. pastoense Baker (1887: 21). Type:—ECUADOR: Without locality, Jameson s. n. (holotype K).
Lycopodium nubigenum Herzog (1915: 2).—Urostachys nubigenus (Herzog) Nessel (1939: 246). Type:—BOLIVIA. La Paz: Comarapa,
T. Herzog 1967 (holotype L; isotypes B, Z).
Illustrations: Øllgaard (1988: Fig. 15B, 17A).
Description:—Slender, pendulous, to 150 cm long, to 10 times dichotomous. Shoots heterophyllous, in the basal
divisions, to 30–100 cm from the base, ca. 10–20(–25) mm in diameter including the expanded leaves, then abruptly
(rarely gradually) constricted to (1–)1.5–2(–2.5) mm in diameter including the imbricate, reduced leaves of the to 50
cm long quadrangular to subterete, terminal divisions. Stems excluding leaves 0.7–1.2 mm thick at the base, upward
tapering to ca. 0.5 mm, greenish to bright red. Expanded leaves of basal divisions borne in alternating whorls of 3, or
decussate, subdecussate or alternate, the pairs or whorls (1.5–)2–6 mm apart, usually widely spaced in alternate-leaved
stem portions, forming 4 or 6 usually indistinct longitudinal ranks, perpendicular to the stem to falcately ascending,
lanceolate to linear-lanceolate, widest at or below the middle, (4–)6–10(–13) × (1–)1.5–2 mm, softly to firmly
herbaceous, with flat to slightly revolute margins, the lamina twisted to a vertical position. Expanded leaves near the
constriction often sporangiate. Leaves of constricted terminal divisions decussate, or subdecussate, continuously or
discontinuously sporangiate, appressed and clasping with their bases, abaxially rounded to carinate, widely lanceolate
to widely ovate or subcordate, acute to mucronate or cuspidate, rarely with an elongate flat apex, 1.7–2(–4) mm long
(in gradually heterophyllous shoots sometimes longer), 1–1.5 mm wide, equalling to more than twice as long as the
sporangia, the leaf pairs (0.5–)1–2(–3) mm apart. Sporangia 1.2–1.5 mm wide.
Distribution:—Costa Rica (?), along the Andes south to northern Argentina, Galápagos Islands.
Habitat:—Epiphytic (rarely rupestral), in upper montane and elfin forest in both cordilleras, 2000–3300 m.
Notes:—Phlegmariurus phylicifolius occurs in a wide range of epiphytic and rupestral habitats with highly
variable exposure and humidity. Its growth habit, i. e. plant length, leaf size, texture and crowding, and stem colour,
responds strongly to the environmental conditions. Two extremes are shown in Øllgaard 1988 (Fig. 15B and Fig.
17A). Forms with a gradual change from expanded to constricted shoots were distinguished by Baker as var. pastoense
(under Lycopodium subulatum), appear to be sporadic aberrants. The gradual transition, and frequent recurrence of
expanded leaves in the otherwise constricted, terminal divisions, is sporadic in several heterophyllous species, and in
one species, P. myrsinites seems to be the common situation.
Phlegmariurus phylicifolius is related to both P. myrsinites, P. subulatus, and to P. biformis (Hooker 1840: t. 228)
B.Øllgaard (2012a: 480), and P. erythrocaulon (Fée 1872–1873: 95) B.Øllgaard (2012a: 480) (both from SE Brazil).
Specimens examined:—Province unknown: Mutis 3363 (F, K, US). Antioquia: Mun. Bello, Vereda San Felix, 8.2
km from Medellín–SanPedro road, 3020 m, Zarucchi et al. 6849 (AAU). Caldas: Summit to Cucarronera, new Quindío
trail, Cord. Central, Killip 9785 (GH, NY, US). Cauca: Puracé, road to parque Nacional de Puracé, 3250 m, Santa &
Hoyos 719 (AAU, COL, HUA, QCA). Páramo de San Pedro, above Totoró, 3000 m, Espinal & Ramos 3507 (COL,
MEDEL). Cundinamarca: La Caldera, 3000 m, Huertas & Camargo (C.M.F.) 12 (F). Sotara, Popoyan, Hartweg 1476
(BR, K, P, US). Usaquén, 2800 m, Schneider 53 (COL, S). Road to Choachí at end of páramo, 3000 m, Acosta-Arteaga
508 (HUA). Salto de Tequendama, Aristé-Joseph A 415 (US). Norte de Santander: Ocaña, 2440–3050 m, Schlim 474
(F, K, P), 475 (BR, F, G, K, P). Putumayo: Valley of Sibundoy, 2000 m, Mora 2109 (US).
Phlegmariurus subulatus (Poiret) B.Øllgaard (2012b: 19).—Fig. 1B.
Lycopodium subulatum Poiret (1814: 544).—Urostachys subulatus (Desvaux ex Poiret) Nessel (1927: 420).—Huperzia subulata (Poiret)
Holub (1985: 77). Type: Herbier de A. N. Desvaux. Donnée par Mme Vve Lavallée en 1896. Annotated by Desvaux (lectotype
P00559228, designated by Øllgaard (1988: 119), possibly a duplicate of a plant in A. Bonpland’s herbarium, bearing the locality:
Quindino, [? Quindió, Colombia], a possible isotype in G.

Urostachys ewanii Herter (1954: 126–127).—Lycopodium ewanii (Herter) Morton (1964: 72). Type:—COLOMBIA: Nariño, Río Potreros,
tributary of Río Tellez, above Soledad, Cord, Or. 3150 m, Ewan 16526 (holotype US; isotype NO).
Illustrations: Øllgaard (1988: Fig. 17B); Lellinger (1989: Fig. 38).
Description:—Slender, pendulous, to 2 m long. Shoots heterophyllous, ca. 15–25 mm in diameter including the
expanded leaves in the basal divisions, abruptly constricted to 1.5–2 mm in diameter including the reduced and
appressed leaves of the terminal divisions. Stems excluding leaves ca. 1 mm thick at the base, upward tapering to less
than 0.5 mm, pale greenish to bright red, at least to 10 times dichotomous. Expanded leaves of basal divisions borne in
alternating, often irregular whorls of 3, or often upward subdecussate, the whorls or leaf pairs 2–4 mm apart, spreading
to perpendicular to the stem, narrowly lanceolate to linear-tapering, straight to obliquely falcate-ascending, (6–)10–15
× 0.5–1 mm, twisted at the base, flat, soft-herbaceous. Leaves of terminal constricted divisions subdecussate, or in
alternating whorls of 3 just above the constriction of the shoots, the leaf pairs or whorls 1–2 mm apart, sporangiate
almost throughout, closely imbricate, widely ovate with obtuse to acute apex, abaxially rounded to carinate, 1.3–2 ×
1.3–1.5 mm, equalling or slightly exceeding the sporangia. Sporangia ca. 1 mm wide.
Distribution:—Honduras, Costa Rica, Panamá, Colombia to Bolivia.
Habitat:—Epiphytic in cloud forest and elfin forest, 2800–3700 m.
Notes:—Closely related to Phlegmariurus phylicifolius with which it has an overlapping distribution, but which occurs
in a wider range of generally more exposed habitats. P. subulatus seems restricted to the most humid forests of the
slopes of the Andes.
Specimens examined:—Antioquia: Mun. Urrao, Jaiperá, Vereda El Chuscal, Páramo de Frontino, Llanogrande,
3000–3390 m, Callejas et al. 7451 (AAU). Caldas: Mountain N of Quebrada Venus, E of Armenia, 3705 m, St.
John 20856 (US). Cauca: Macizo Colombiano, Páramo de las Papas, around Laguna Cusiyaco, 3470 m, Barclay &
Juajibioy 5967 (GH, MO). Cord. Central, W slope, headwaters of Río Palo, quebrada del Río López, Quebrada del
Duende, 3400–3450 m, Cuatrecasas 18928 (F, GH, S, US). Macizo Colombiano, Páramo de las Papas, around Laguna
Cusiyaco, 3200 m, Idrobo et al. 4007 (COL, NY). Paletará, Cord. Centr., forest E of Llano, 3100–3300 m, Pennell
7019 (GH, NY, US). Páramo de Buena Vista, Huila Group, Centr, Cord. 3200 m, Pittier 1193 (US). Cord. Central,
Puracé, 3450 m, Sneidern 1697 (GH, NY, S). Paletará, 30 km SE of Popoyan, 3100 m, Tryon & Tryon 5981 (COL, GH,
P, US). Road Totoró–Inzá, between Páramo de Las Delicias and Inzá, E slopes of Cord. Central, 3100 m, Santa et al.
981 (AAU). Road Totoró–Inzá, ascending to Páramo de Las Delicias, E slopes of Cord. Central, 3090 m, Santa et al.
973 (AAU). Cundinamarca: Montserrat, Lindig s.n. (US). Huila/Cauca: Macizo Colombiano, Páramo de las Papas,
around Laguna La Magdalena, 3530 m, Idrobo et al. 2995 (COL, G, NY), 3320 (COL, G, VEN). Nariño: Páramo de
Tábano, alto de Cord. between Pasto and El Encano, W slope, 3200 m, Cuatrecasas 11927 (COL, F, HUA, UC, US).
Drainage of upper Río Tescual, above Cordoba, 2840–2890 m, Ewan 16293 (GH, NO, US). Pasto:1843, Jameson 424
(E, G, K, NY frag., US). Risaralda/Valle del Cauca: Cerro Tatama, Cord. Occidenal, 3200–3400 m, Pennell 10492
(GH, US). Tolima: Mariquita, versant du Tolima, 1200–1300 toises, Linden 1001 (P).
Group of Phlegmariurus acerosus
Phlegmariurus acerosus (Swartz) B.Øllgaard (2012a: 480).—Fig. 7A.
Lycopodium acerosum Swartz (1806b: 1575).—Urostachys acerosus (Swartz) Nessel (1927: 399).—Huperzia acerosa (Swartz) Holub
(1985: 70).—Lectotype (designated by Proctor (1977: 25): Plumier (1705: t. 166B); an authentic specimen possibly in Herb. Suriani
no. 635 (P, not seen).
Illustrations: Øllgaard (1988: Fig. 18C).
Description:—Very delicate plants, flaccidly pendulous, at least to 70 cm long. Shoots usually gradually heterophyllous,
sometimes homophyllous, 3–6 mm in diameter including the leaves in basal divisions, tapering to 1–3(–6) mm in
terminal divisions. Stems excluding leaves 0.6–1 mm thick at the base, tapering to ca. 0.3–0.5 mm upward, somewhat
concealed by the leaves, pale greenish to stramineous, at least to 10 times dichotomous, usually densely, unilaterally
or omnilaterally sporangiate in separate, periodically produced zones of the terminal divisions, or continuously
sporangiate from 15–30 cm above the base and upward. Leaves gradually modified along the stems. Leaves of basal
divisions densely crowded, borne in irregular alternating whorls of 6–7, these 0.5–2 mm apart, forming 12–14 indistinct
longitudinal ranks, ascending and upward curved to appressed or somewhat secund, acicular-filiform, narrowly and
prominently decurrent, 3.5–5 × 0.2–0.4 mm, soft herbaceous to subcoriaceous, convex below, canaliculate above, with
involute margins. Vegetative leaves of terminal constricted divisions borne in irregular whorls of 4–6, these 0.5–2 mm
apart, forming 8–12 indistinct longitudinal ranks, ascending to closely appressed, acicular-filiform to linear-lanceolate,
2–4 mm long, otherwise conform. Sporophylls borne in irregular, alternating whorls of 3–5, conform, or shorter
and wider, linear-lanceolate and long acuminate to lanceolate, usually appressed, 1.5–4 × 0.3–0.8 mm, rounded to
subcarinate abaxially. Sporangia 0.7–1 mm wide.
Distribution:—West Indies, Guatemala, Costa Rica, northern South America, south to Ecuador, Bolivia, northern
Argentina and southeastern Brazil.
Habitat:—Epiphytic, in lowland wet forest, elev. 500–2100 m.
FIGURE 7. A. Phlegmariurus acerosus (Holdridge 1652, Ecuador, US) growth habit. A1. Close-up of basal division. A2. Close-up
of terminal sporangiate division.—B. Phlegmariurus filiformis (Ecuador, Espinosa E-1490, GH), growth habit. B1. Close-up of basal
division. B2. Close-up of terminal sporangiate division, B3. Close-up of terminal sporangiate division. Del K. Tind.

Notes:—In older literature Phlegmariurus acerosus has often been referred to P. verticillatus (Linnaeus f. 1882:
448) Field & al. (2015: 21), or Lycopodium setaceum Lamarck (1792: 563), both of which are based on material from
Réunion Island, and belong to a paleotropical clade (Wikström & Kenrick 2000, Field et al. 2015). The plants from the
Old World referred to Phlegmariurus verticillatus resemble P. acerosus, but differ from this in a more robust growth
habit, usually longer and more crowded leaves, and usually more densely and omnilaterally sporangiate terminal
divisions. For discussion of related species see P. curvifolius.
This is the most widespread and apparently least specialized of the species. The name Lycopodium portoricense
Underw. & Lloyd (1906: 108) represents some West Indian plants, relatively coarse and vigorous, that are usually
terrestrial, hanging over banks. They are closely related to, or perhaps conspecific with Phlegmariurus acerosus.
Specimens examined:—Province unknown: Lehmann 4525 (US). Valle del Cauca: La Cumbre, 1600–2100 m,
Killip 11609 (GH, NY, US).
Phlegmariurus curvifolius (Kunze) B.Øllgaard (2012b: 14).—Fig. 8C.
Lycopodium curvifolium Kunze (1835: 5).—Lycopodium tenue Willdenow var. tenuissimum Spring (1849: 21).—Urostachys curvifolius
(Kunze) Nessel (1939: 129).—Huperzia curvifolia (Kunze) Holub (1985: 72). Type:—PERU. Huánuco: “sylvar. densarum arboribus
prope Pampayaco”, 1829, Poeppig s.n. (AWH, B, BR, E, G, K, M, NY, W).
Lycopodium cernuum Linnaeus (1753: 1103) var. tenerrimum Nessel (1940a: 173). Type:—ECUADOR: Without locality, Sodiro n. 3101”
(BONN-Nessel 535).
Illustrations: Øllgaard (1988: Fig. 20C)
Description:—Delicate to extremely delicate plants, flaccidly pendulous, to 60 cm long. Shoots gradually

heterophyllous, 2.5–6 mm in diameter including the leaves in basal divisions, tapering to 0.3–1.5(–2) mm in diameter
including leaves in terminal divisions. Stems excluding leaves 0.4–0.7 mm thick at the base, tapering to 0.2–0.4
mm upward, greenish to stramineous, usually not concealed by leaves, to more than 10 times dichotomous, usually
sparsely sporangiate from 10–20 cm above the base and upward. Leaves of basal divisions borne in alternating whorls
of 4–5, these 1–2 mm apart, forming 8–10 indistinct longitudinal ranks, patently sigmoid to strongly falcately upward
curved, acicular-filiform, long and widely decurrent, 2–4(–5) × 0.2–0.3 mm, extremely delicate, abaxially convex, flat
or canaliculate above. Vegetative leaves of terminal constricted divisions alternate, or borne in alternating irregular
whorls of 3, closely appressed, lanceolate to linear-lanceolate, somewhat clasping with the base, 1.2–3.5 × 0.2–0.6
mm, usually strongly involute, often sharply carinate at the base, short to long decurrent. Sporangia usually few, borne,
often unilaterally, in the axils of every third to fifteenth (or more) leaf along the terminal shoots. Sporophylls conform,
or shorter and wider, lanceolate to widely ovate, or subcordate and short to long acuminate, usually widely spreading,
1–3 × 0.3–0.8 mm, in ultimate divisions sometimes scarcely exceeding the sporangia, abaxially with a prominent vein,
or carinate. Sporangia 0.6–0.8 mm wide.
Distribution:—Costa Rica to Peru.
Habitat:—Epiphytic in lower, wet montane forest, elev. 150–500 m.
Notes:—Phlegmariurus curvifolius is variable in size and compactness. This variation may probably not be
explained by environmental factors alone. The species is easily recognized by the strongly upward curved (falcate)
leaves of basal divisions, and the appressed non-sporangiate leaves of the narrow, terminal divisions. The smallest forms
with hair-like terminal shoots represent the most fragile and slender extreme known in the family. They correspond to
the type collection. Specimens examined:—Cauca: Near Río Timbiquí, 200–500 m, Lehmann 8909 (K, US). Chocó:
5–8 km E of Playa de Oro (E of Tado), 140 m, Juncosa 2510 (MO).
Phlegmariurus filiformis (Swartz) W.H. Wagner (1995: 242).—Fig. 7B.
Lycopodium filiforme Swartz (1801: 114).—Urostachys filiformis (Swartz) Herter (1949: 61).—Lycopodium acerosum Swartz var filiforme
(Swartz) Spring (1840: 111).—Lycopodium verticillatum L. f. (1782: 448) var. filiforme (Swartz) Spring (1842: 47).—Huperzia
verticillata (Linnaeus f.) Trevisan var. filiformis (Swartz) Trevisan (1874: 248).—Huperzia filiformis (Swartz) Holub (1985: 72).
Type:—USA. Hawai‘i: “Insulae Sandwich Oceani pacifici”, specimen not located (S?).
Lycopodium polytrichoides Kaulfuss (1824: 6).—Huperzia polytrichoides (Kaulfuss) Trevisan (1874: 248).—Urostachys polytrichoides
(Kaulfuss) Nessel (1939: 129). Type:—USA. Hawai‘i: OWahu insularum Sandwich, Chamisso s. n. (not located, Leipzig, destroyed
?).
Illustrations: Lellinger (1989: Fig. 40); Øllgaard (1988: Fig. 21A).
Description:—Slender, flaccidly pendulous plants, at least to 45 cm long. Shoots gradually heterophyllous, 4–5(–6)
mm in diameter including the leaves in basal divisions, tapering to 1–2 mm in terminal divisions. Stems excluding
leaves 0.5–1 mm thick at the base, tapering to ca. 0.5 mm upward, almost completely concealed by the leaves, pale
greenish to stramineous, to 9 times dichotomous, usually densely, unilaterally to omnilaterally sporangiate in short
zones of the terminal divisions from 7–30 cm above the base and upward. Leaves gradually modified along the stems.
Leaves of basal divisions densely crowded, borne in irregular alternating whorls of 4–6, forming 8–12 indistinct
longitudinal ranks, ascending to closely imbricate or somewhat secund, acicular to linear, narrowly decurrent, 4–5.5 ×
0.2–0.4 mm wide just above the widened base, soft herbaceous to coriaceous, convex below, canaliculate above, with
involute margins. Vegetative leaves of terminal constricted divisions borne in irregular whorls of 3–4, these 1–2 mm
apart, forming 6–8 indistinct longitudinal ranks, closely appressed, lanceolate to linear-lanceolate, 1.3–3 × 0.4–0.6
mm, usually strongly involute (dried), short to long decurrent. Sporophylls conform, or shorter and wider, lanceolate
and long acuminate to widely ovate to subcordate and acute or acuminate, usually appressed, 1–2.5 × 0.4–0.8 mm, in
ultimate divisions equalling to at least twice as long as the sporangia, rounded to carinate abaxially. Sporangia 0.8–1.2
mm wide.
Distribution:—Hawaiian Islands, Central America, Andes from Venezuela to Bolivia.
Habitat:—Epiphytic, in lower montane forest, 500–1500 m.
Notes:—Phlegmariurus filiformis is closely related to P. acerosus and P. curvifolius, and their delimitation is
sometimes problematic. It appears to be adapted to slightly drier growth conditions than P. acerosus.
Specimens examined:—César: Pueblo Bello, Sierra Nevada de Santa Marta, 1220 m, Foster & Smith 1495 (A,
COL). San Antonio, Sierra de Santa Marta, Hanbury-Tracy 556 (K). Meta: San Juan de Arama, left bank of Río Güejar
near landing strip Los Micos, 500 m, Idrobo & Schultes 1262 (COL, GH, US).
Phlegmariurus tenuis (Willdenow) B.Øllgaard (2012b: 19).—Fig. 8A–B.
Lycopodium tenue Willdenow (1810: 55).—Huperzia tenuis (Willdenow) Trevisan (1874: 248).—Urostachys tenuis (Willdenow) Nessel
(1927: 401). Type:—ECUADOR. Loja: Valley of Vinajacu, near Loja, Humboldt & Bonpland 3363 (holotype B-Willdenow no.
19423; isotypes, BM, LG, P00669229).
Illustrations: Øllgaard (1988: Fig. 20A, B).
Description:—Usually epiphytic, delicate to extremely delicate, flaccidly pendulous, at least to 75 cm long. Shoots
gradually heterophyllous, 3–8 mm in diameter including the leaves in the basal divisions, tapering to 1–3 mm in
diameter in the narrow terminal divisions. Stems excluding leaves 0.5–0.7 mm thick at the base, tapering to ca. 0.3
mm upward, pale greenish to stramineous, at least to 10 times dichotomous, usually discontinuously and unilaterally
sporangiate from 10–30 cm above the base and upward. Leaves of basal divisions borne in irregular alternating
whorls of 3–5, or spirally arranged, forming 6–10 indistinct longitudinal ranks, usually densely crowded, spreading
to ascending, straight to somewhat curved, often secund, acicular, tapering from a subauriculate base, 3–5(–7) ×
0.3–0.5 mm, decurrent. Vegetative leaves and sporophylls of constricted terminal divisions almost conform, borne in
alternating irregular whorls of 3, or irregularly alternate, patently diverging from the stem, with upward curved tips,
widely ovate to subhastate, 1–3 × 0.5–1.4 mm wide at the base, clasping with the base, tapering to abruptly narrowed
into a short to long acuminate apex, bluntly to sharply carinate, usually unilaterally, discontinuously, but often densely
sporangiate. Sporangia 0.8–1.2 mm wide.
Distribution:—Costa Rica, Panama, Andes from Venezuela to Bolivia
Habitat:—Epiphytic in uppermost montane forests, elev. 3000–3500 m.
Notes:—Phlegmariurus tenuis exhibits the same type of size variation as mentioned for P. curvifolius. Some
forms are almost as extremely fragile and slender, but are easily distinguished by the spreading leaves in the terminal
divisions. This species is also ecologically distinct, being restricted to the uppermost montane forests.
Specimens examined:—Cauca: Macizo Colombiano, páramo de Las Papas, Boqueron–La Hoyola, 3200–3510
m, Idrobo et al. 3446 (COL, G). Nariño: Near Pasto, Jameson 423 (BM, E, G, GH, OXF, US). Between Pususquer and
Piedra Aucha, Lehmann 5021 (AAU, F, GH, K, US). Norte de Santander: Ocaña, 3050–3350 m, Schlim 342 (BR, G,
P).

FIGURE 8. A. Phlegmariurus tenuis (Spruce 5604, E) growth habit. A1. Close-up of basal division. A2. Close-up of terminal sporangiate
division.—B. Phlegmariurus tenuis (Ecuador: Holm-Nielsen et al. 5342, AAU) growth habit. B1. Close-up of basal division. B2. Close-
up of terminal sporangiate division.—C. Phlegmariurus curvifolius composite (Lugo 848, AAU) Growth habit. C1. (Ecuador: Harling
& Andersson 17354, AAU) close-up of basal division. C2. (Ecuador, Lugo 848, AAU) close-up of terminal sporangiate division. Del. B.
Johnsen.
Group of Phlegmariurus dichotomus
Phlegmariurus arcuatus (B.Øllg.) B.Øllgaard (2012b: 12).—Fig. 9A.
Huperzia arcuata B.Øllgaard (1988: 15). Type:—ECUADOR. Carchi: Road El Angel–Tulcán, 3500 m, Holm-Nielsen, Jeppesen, Løjtnant
& Øllgaard 5341 (holotype AAU; isotypes GB, QCA, US).
Illustrations: Øllgaard (1988: Fig. 1D)
FIGURE 9. A. Phlegmariurus arcuatus (Ecuador: Holm-Nielsen et al. 5341, AAU) growth habit. A1. Close-up of sporangiate division.
Del B. Johnsen.—B. Phlegmariurus brongniartii (Maas & Plowman 2121, AAU) growth habit. B1 Close-up of sporangiate division. B2.
Close-up of sporophyll. Del K. Tind.
Description:—Plants ascending to erect from a decumbent base, with nodding shoot apices, to 30 cm tall (terrestrial),
or recurved to pendulous and to 90 cm long (epiphytes), sparsely branched, to 5 times dichotomous. Shoots
homophyllous, equally thick throughout, 25–30 mm in diameter, or tapering to ca. 12 mm in diameter including leaves.
Stem excluding leaves 2–3 mm thick at base, tapering to ca. 1.5 mm upward, sporangiate throughout from near the
base, or with separate sporangiate zones along terminal divisions. Leaves borne in more or less regular, often oblique,

alternating whorls of 6–7 near the base, upward of 5–6, these 2–3 mm apart, spreading to falcately ascending, linear to
subulate from a slightly widened lamina base, (10–)14–19 × (0.5–)0.7–1 mm wide above the widened base, gradually
tapering from the middle or below the middle, adaxially flat–slightly concave, or with slightly revolute margins, with
abaxially slightly prominent vein and smooth margins. Lamina base slightly widened, 1–1.2 mm wide, twisting the
lamina to a vertical position. Vegetative leaves of basal divisions often spreading to reflexed and not twisted. Sporangia
1.3–2 mm wide.
Distribution:—Southern Colombia, Ecuador, Peru (Amazonas, San Martín).
Habitat:—Upper montane forest, near the timber line, on the forest floor, in semishade, or epiphytic, elev. 3000–
3600 m. Ramírez Padilla & Macias Pinto (2007) reported this species from Dept. Cauca.
Notes:—Closely related to Phlegmariurus hippurideus, from which it differs by its obliquely falcate-ascending,
twisted leaves, and the nodding to pendulous shoot tips. Phlegmariurus arcuatus also resembles P. dichotomus, but is
larger in all parts, and occurs mainly in timber line forest habitats.
Specimens examined:—Nariño: Páramo del Tábano, Pasto–el Encano, W slope, 3200 m, Cuatrecasas 11926
(COL, US). Páramo del Tábano, Pasto–el Encano, W slope, 3200 m, André, K 416 (NY). Risaralda: Parque Los
Nevados, near Finca El Bosque, 3600 m, Verweij & Beukema F 15aa (U).
26. Phlegmariurus dichotomus (Jaquin) W.H.Wagner (1993: 305).—Fig. 5B.
Lycopodium dichotomum Jacquin (1762: 314).—Huperzia dichotoma (Jacquin) Trevisan (1874: 248).—Urostachys dichotomus (Jacquin)
Herter (1922: 249). Type:—Fide Proctor (1977: 28): Jacquin (1776: t. 45) lectotype, an authentic specimen in BM (Exemplar, quod
collegi in insula Martinica, et quod iconi inservitt mihi olim).
Lycopodium gramineum Spring (1849: 19).—Urostachys gramineus (Spring) Herter (1949: 63). Type:—ECUADOR. Guayaquil: Jameson
s. n. (holotype K).
Lycopodium barbatum Christ (1905: 254); non Kaulfuss (1824: Selaginella).—Urostachys barbatus (Christ) Nessel (1939: 30). Type:—
COSTA RICA: Werclé s. n. (holotype P [Herb. Christ]; isotype BONN [Herb. Nessel 7, fragment]).
Lycopodium chamaepeuce Herter (1909a: 50).—Urostachys chamaepeuce (Herter) Herter (1923: 164). Type: None designated.
Urostachys chamaepeuce (Herter) Herter var. urbanianus Herter (1923: 164). Type:—GRENADA: Annandale, February 1906, Broadway
3457 (B n.v.; NY), 3458b (B n.v.).
Urostachys lindeneri Nessel (1940a: 164 excluding figure).—Huperzia lindeneri (Nessel) Holub (1985: 74). Type:—GUATEMALA:
Chilion, Mazatenango, Bernoulli & Cario s. n. (holotype BONN-Nessel 350).
Urostachys schlechtendalii Nessel (1940a: 153).—Huperzia schlechtendalii (Nessel) Holub (1985: 76). Type:—MEXICO: “Guaveyma”,
Wagner s.n. (holotype BONN-Nessel 349 p. p.).
Illustrations: Lellinger (1989: Fig. 23); Øllgaard (1988: Fig. 19D).
Description:—Plants pendent or recurved from an erect base, lax to subrigid, sparsely to densely and spreadingly
branched, usually to 25(–50) cm long. Shoots homophyllous, almost equally thick throughout, 20–25 mm in diameter
including leaves near the base, usually tapering to (10–)15 mm in densely sporangiate terminal divisions. Stem
excluding leaves 2–3 mm thick at the base, sometimes tapering to 1.5–2 mm, usually almost concealed by leaves,
ridged by decurrent leaf bases, sporangiate from 5–15 cm above the base and upward, usually 3–5 times dichotomous.
Leaves borne in alternating whorls of 5–6, these 1–2 mm apart, forming 10–12 indistinct longitudinal ranks, spreading
to ascending, usually somewhat obliquely falcate, in terminal divisions ascending to somewhat appressed, or appressed
throughout, linear-subulate, gradually tapering from the base, (8–)10–15(–17) × 0.7–1 mm, almost flat, with slightly
prominent vein above, with flat or slightly revolute margins, abaxially with slightly to sharply prominent vein, soft-
herbaceous, usually twisted at the base. Leaf bases long and prominently decurrent, green, as wide as, or wider than
the lamina, with sharply prominent vein. Sporangia 1–1.5 mm wide.
Distribution:—Florida, Mesoamerica, West Indies, Colombia, Venezuela, Ecuador, Galapagos Islands, Peru,
Bolivia, Brazil.
Habitat:—Usually epiphytic in lowland, riverine, and premontane forests. Elev. 0–1400 m.
Notes:—Plants usually rather short and recurved to pendulous, with linear, densely crowded, usually basally
twisted, falcately ascending leaves. This is the most widespread and variable species of the group. Closer study of the
material referred to this species may show the presence of more than one taxon, especially in the northern part of the
range, because there is considerable variation in the thickness of stems, leaf length and direction, and compactness of
the plants. However, the correlation of the variation with growth conditions is uncertain at this point.
Urostachys mortonii Herter (1930: 108).—Huperzia mortonii (Herter) Holub (1985: 75).—Syntypes from
Guatemala, n. v., seems to be a synonym of P. dichotomus.
Specimens examined:—Antioquia: Valle del Río Anorí, between Dos Bocas and Anorí, Providencia, 400–900
m, Fonnegra et al. 320 (HUA). Cauca: Pacific coast, Río Micay in Guayabal, 5–20 m, Cuatrecasas 14159A (US).
SW of El Tambo, Quebrada La Pita SW of Tambo, 1100 m, Haught 5270 (COL, US). El Tambo, hoya del Patía,
Corregimiento de Mosquera, 1100 m, Idrobo & Fernández 152 (COL, US). Chocó: Río Calima (Chocó), La Trojita, 5–
50 m, Cuatrecasas 16506 (S, US). Drainage of Río San Juan, near Palestina, 5 m, Forero et al. 3792 (MO). Magdalena:
Santa Marta, near Valparaiso, 1070 m, Smith 2237 (F, K, MO, NY, P, S, US). Valle del Cauca: Mun. Zarzal, road La
Paila–Zarzal, Hda. El Medio, 975 m, Silverstone-Sopkin & Paz 3135 (AAU).
Phlegmariurus hippurideus (Christ) B.Øllgaard (2012: 480).—Fig. 10A-B.
Lycopodium hippurideum Christ (1901: 56).—Urostachys hippurideus (Christ) Nessel (1939: 88).—Huperzia hippuridea (Christ) Holub
(1985: 73). Type:—COSTA RICA. San José: El Páramo, massif de Buena Vista, 3000 m, 1897, Pittier 10619 (holotype P00559179;
isotype US).
Lycopodium lechleri Hieronymus (1905: 571).—Urostachys lechleri (Hieronymus) Nessel (1939: 85).—Huperzia lechleri (Hieronymus)
Holub (1985: 74). Type:—PERU: Tabina, Lechler, ed. Hohenacker 2012 (lectotype B, designated by Rolleri (1981: 82); isolectotypes
G, K, P, UPS).
Urostachys lechleri Hieronymus var. lehmannii Nessel (1940a: 161).—Urostachys lehmannii (Nessel) Herter (1949: 67).—Syntypes:
BONN, Herb. Nessel 148, two pairs of fragments indicated to be from Ecuador (Spruce s.n. Pallatanga 1858, marked 7; the other
is indicated to be from Ecuador: unknown collector s.n. »Aus dem Herbar Bonaparte, Paris«, annotated »Ur. lehmannii Boge.«). A
different specimen from Bolivia (Mandon 1588) is illustrated in t. 8, f. 35 (l. c.).
Urostachys poseidonis Herter (1954: 122–123).—Lycopodium poseidonis (Herter) Morton, C. (1964: 42). Type:—ECUADOR.
Chimborazo: Peripa, “In regione arborea superiore prope pagum Peripe”, 3400 m, December 1921, Rose, in Mille 35 (holotype US,
isotypes GH, NY).
Lycopodium bolivianum Rosenstock (1912: 59).—Urostachys bolivianus (Rosenstock) Nessel (1927: 396).—Huperzia boliviana
(Rosenstock) Rolleri & Deferrari (1988: 155). Type:—BOLIVIA. La Paz: [Prov. Nor Yungas] Unduavi, Buchtien 2632 (isotypes L,
Z).
Illustrations: Øllgaard (1988: Fig. 1A, 2A.); Mickel & Beitel (1988: Fig. 2E); Lellinger (1989: Fig. 27).
Description:—Plants ascending to stiffly erect from a decumbent base, to 60 cm tall, sparsely branched, to 3(–5) times
dichotomous. Shoots homophyllous, equally thick throughout, 10–35 mm in diameter including leaves, sporangiate
from ca. 20 cm above the ground, often in separate zones. Stems excluding leaves 2.5–4 mm thick at the base, tapering
to ca. 2–3 mm upward. Leaves borne in more or less regular, often oblique, alternating whorls of 5–8, these 2–5 mm
apart, forming 10–16 indistinct longitudinal ranks, spreading to reflexed, sometimes sharply reflexed and appressed to
the stem, linear to linear-subulate, evenly tapering from the base or the middle, (10–)11–19 × 0.8–1.3 mm wide at the
base, not, or rarely, twisted at base, broadly decurrent, adaxially slightly convex–slightly concave or canaliculate, with
slightly prominent vein, abaxially with slightly prominent vein, with smooth, sometimes slightly revolute margins,
with indistinctly to prominently and broadly decurrent leaf bases. Sporangia 1.5–2 mm wide.
Distribution:—Central America, Greater Antilles, Andes from Venezuela south to Bolivia, the Venezuelan and
Brazilian Roraima Formation.
Habitat:—Upper montane forest, especially near the forest limit, usually on the forest floor, rarely epiphytic, in
semishade, elev. 2200–3750 m.
Notes:—Phlegmariurus hippurideus belongs to a group of closely related taxa of high montane forests throughout
tropical America, including also P. arcuatus, and the Brazilian P. nudus (Nessel) B.Øllgaard (2012a: 481).
Specimens examined:—Antioquia: Mun. San Cristóbal, paraje Boquerón, along road to the sea, 2200–2700
m, Hernández et al. 531 (HUA). Mun. Urrao, Encarnación, 1 hour S of Páramo de Frontino, El Río, 3000–3150 m,
Callejas et al. 7833 (AAU). Antioquia, Kalbreyer 454 (K). Arauca: Sierra Nevada del Cocuy, Quebrada El Playón,
3000 m, Cleef 10146 (AAU, COL, U). Boyacá: Sierra Nevada del Cocuy, around Cobugón, 2900 m, Grubb & Guymer
P 51 (COL, US). Caldas: Cerro Tatama, Cordillera Occidental,, 2800–3300 m, Pennell 10433 (US). Chilí páramo,
3750 m, St.John 20891 (US). Cauca: Macizo Colombiano, páramo de las Papas, around Laguna de Cusiyaco, 3450 m,
Barclay & Juajibioy 6005 (GH). Cord. Central, W slope, headwaters of Río Palo, Quebrada de Santo Domingo, 2950–
3150 m, Cuatrecasas 19237 (F, GH, US). Macizo Colombiano, páramo de las Papas, near Laguna La Magdalena,
3530 m, Idrobo et al. 2944 (COL, G). Paletará, Cord. Central, 3100–3300 m, Pennell 7014 (GH, NY, US). Mt. Santa
Ana, cord. occ. 2700–3000 m, Pennell 7471 (GH, NY, US). Cundinamarca: Usaquén, Amórtegui A-40 (US). Bogotá,
Aristé-Joseph 290 (US). Macizo de Bogotá, Quebrada del Rosal, 3200 m, Cuatrecasas 5718 (COL, US). Zipaquirá,
Páramo de Guerrero, 2800–3000 m, García-Barriga 20783 (GH, US). Cord. Or., Cerro Negro, Hato Grande, ridge E
of Río Muchindote, 15 km E of Cachetá, 2870 m, Grant 9409 (US). Mun. Fomeque, Páramo de Chingaza, La Laja,
2700–3250 m, Huertas & Camargo 5777 (COL). Usaquén, 2700 m, Schneider 386 (S). Páramo de Chipaque, 3100–
3200 m, Schultes 1047 (GH, US). Bojacá, Bosques de La Merced, 2970 m, Torres 107 (COL). César: Sierra de Perijá,
E of Manaure, Quebrada de Floridablanca, 2700–2800 m, Cuatrecasas & Romero Castañeda 25247 (COL, US). Sierra
de Perijá, 11 km ENE of Manaure, 47 km E of Valledupar, 2 km from the Venezuelan border, 2700 m, Grant 10839
(US). Nariño: Km 13–14 W of Pasto towards Laguna La Cocha, 3230 m, Luteyn et al. 5009 (AAU). Drainage of upper
Río Tescual, above Cordoba, 2840–2890 m, Ewan 16295 (NO, UC, US). Norte de Santander: Pamplona, Las Cruces,
3050 m, Funck & Schlim 1473 (BR, G, P). E Cordillera, Páramo de Hatico, on road Toledo–Pamplona, 2900 m, Killip
& Smith 20649 (GH, NY, US). Ocaña, 2440–3050 m, Schlim 471 (G), 473 (BR, G, P). Putumayo: W side of Laguna
de La Cocha, 5 km S of El Encano, 2900–3000 m, Ewan 16334 (NO, US). Risaralda: Mun. Santuario, NP Tatamá,
2800–3000 m, Stancik 3424 (AAU). Tolima: Quindiu road, 3000 m, Alston 7776 (BM).
FIGURE 10. A. Phlegmariurus hippurideus, (Asplund 9812, Ecuador, S) growth habit. A1, close-up of sporangiate division.—B.
Phlegmariurus hippurideus (Sneidern 2196, Ecuador, S) growth habit, B1. Close-up of sporangiate division.—C. Phlegmariurus urbanii
(Ecuador Asplund 16836, S) growth habit. C1. Close-up of sporangiate division.—D. Phlegmariurus firmus (Tryon & Tryon 5973, NY).
D1. Close-up of sporangiate division. Del. B. Johnsen
Phlegmariurus mollicomus (Spring) B.Øllgaard (2012a: 481).—Fig. 11.
Lycopodium dichotomum Jacquin ssp. mollicomum Martius ex Spring (1838: 162).—Lycopodium mollicomum Spring (1840: 113).—
Lycopodium setaceum Hamilton var. brasiliense Spring (1842: 43).—Huperzia setacea (Hamilton ex D. Don 1825: 18) Trevisan var.
mollicoma (Spring) Trevisan (1874: 248).—Urostachys mollicomus (Spring) Nessel (1927: 407).—Urostachys setaceus (Hamilton)
Nessel var. brasiliensis (Spring) Nessel (1927: 408).—Huperzia mollicoma (Spring) Holub (1985: 75). Type:—BRAZIL: Pará;
“Sylvis Paraënsis”, Martius s. n. (holotype M).
Lycopodium flaccidum Fée (1872–73: 92), non Bory (1833: 9) = Selaginella.—Urostachys flaccidus (Fée) Herter (1923: 164).—Huperzia
flaccida (Fée) Holub (1985: 72).—Urostachys neptuni Herter (1949: 72). Type:—BRAZIL. Rio de Janeiro: “Brasilia fluminensi,
Alto Macahe”, Glaziou 4677 (holotype P; isotypes B, C, RB).
Lycopodium williamsii Underwood & F.Lloyd (1906: 112).—Urostachys williamsii (Underwood & Lloyd) Nessel (1939: 164). Type:—
BOLIVIA: New Brazil, 5,500 ft ., Williams 1393 (holotype NY; isotypes AAU, GH, US).
Illustrations: Øllgaard (1988: Fig. 18B); Lellinger (1989: Fig. 31).
FIGURE 11. Phlegmariurus mollicomus (Plowman & Martinelli 10128, Brazil, AAU) growth habit. A1. Close-up of sporangiate
divisions.
Description:—Pendulous plants, usually with flaccidly hanging divisions, to 30 cm long. Shoots homophyllous or
almost so, almost equally thick throughout, 7–15 mm in diameter including the leaves, or gradually tapering to ca. 5
mm in diameter. Stems excluding leaves 0.5–1 mm thick at the base, often tapering to 0.3 mm, prominently to sharply

ridged by decurrent leaf veins, discontinuously or ultimately continuously sporangiate from 6–15 cm above the base
and upward, to 6(–8) times dichotomous. Leaves borne in alternating, irregular, often oblique whorls of 4–5 in basal
divisions, or of 3–4 in terminal divisions, the whorls 1–2 mm apart, forming 6–10 obscure longitudinal ranks, in basal
divisions ascending to patent, upward more appressed, rarely slightly twisted at the base, straight to slightly upward
curved, linear to linear-subulate. Leaves of basal divisions (7–)10–13 × 0.5–0.7 mm, not or very slightly widened at
the base, adaxially flat or convex, usually somewhat revolute (dried), abaxially with sharply prominent vein at least in
the basal half. Leaves of terminal sporangiate divisions 7–11 mm long, usually loosely appressed, otherwise conform,
or almost flat, with less prominent vein, often slightly widened at the base (not auriculate), rarely tinged with red on
the margins near the base. Decurrent leaf bases not wider than lamina. Sporangia ca. 1 mm wide.
Distribution:—Hispaniola, Costa Rica, Panamá, Colombia, Venezuela, central and southeastern Brazil, Ecuador,
Bolivia. Ramírez Padilla & Macias Pinto (2007) reported this species from Dept. Cauca.
Habitat:—Epiphytic in lower montane forests, elev. ca. 700–1700 m.
Notes:—Phlegmariurus mollicomus is rather similar to P. sarmentosus and P. watsonianus (Maxon 1912: 3)
B.Øllgaard (2012b: 20). It differs from both in the lack of leaf base auricles, and in the sharply prominent leaf vein and
revolute margins. Molecular data (Testo et al. 2018) point to a closer relationship to the group of P. dichotomus and P.
hippurideus.
Specimens examined:—Antioquia: Kalbreyer 1425 (K). Chocó: Km 137 from Caldas on road to Quibdó, 900
m, Madison 1377 (GH). Santander: Charalá, Virolín, trail to El Reloj, 1600–1700 m, Díaz 1535 (COL).
Phlegmariurus pithyoides (Schlechtendal & Chamisso) B.Øllgaard (2012b: 17).—Fig. 12B.
Lycopodium pithyoides Schlechtendal & Chamisso (1830: 623).—Lycopodium mandiocanum Raddi (1819: 280) var. mexicanum Spring
(1842:45).—Urostachys pithyoides (Schlechtendal & Chamisso) Herter (1923: 164).—Huperzia pithyoides (Schlechtendal &
Chamisso) Holub (1985: 76). Type:—MEXICO: Schiede & Deppe s.n. (holotype B; possible isotype HBG [Deppe 832]).
Lycopodium gigas Herter (1909a: 50).—Urostachys gigas (Herter) Herter (1923: 164). Type:—MEXICO: “Südmexico. Epiphyt auf
Bäumen, besonders Eichen, Hänge der Cordilleren”, 2000 m, Haarström s.n. (lectotype P, designated by Herter 1923: 164).
Illustrations: Øllgaard (1992: Fig. 2); Mickel & Beitel (1988: Fig. 1C, D); Lellinger (1989: Fig. 34).
Description:—Erect (or rarely recurved to pendent with age), usually to 25 cm tall (to 35 cm long) 3–4(–8) times
dichotomous, terminal divisions of old plants sometimes densely aggregated. Shoots equally thick throughout, 4–7 cm
in diameter, or tapering to ca. 2 cm in old plants. Stems excluding leaves 4–6 mm thick at base, distally tapering to 2–3
mm, usually completely covered by the wide, bright red, decurrent leaf bases. Leaves uniform throughout, or gradually
smaller distally, borne in irregular alternating whorls of 4–5, spreading to reflexed, usually straight, sometimes with
a basal twist and falcately ascending in distal divisions, linear, tapering in the apical half, (10–)20–35(–45) × 0.7–1.2
mm, bisulcate above (dried), with prominently tumid vein and margins, somewhat coriaceous, the vein prominent to
sunken beneath, margins entire, revolute. Leaf bases with prominently decurrent margins, and a median veinal ridge,
widening to ca. 2 mm. Sporangia ca. 2 mm wide.
Distribution:—Southern Mexico to Costa Rica, Greater Antilles, Colombia, Venezuela (Falcón).
Habitat:—Epiphytic, often on oak trees, in lower to mid-altitude montane rain forests, or seasonal evergreen
forests, elev. 2000 m.
Notes:—One of the most impressive Phlegmariurus species, with the aspect of a small pine tree, with thick,
bright red stems due to the color of the prominent, decurrent leaf bases, and spreading, coriaceous, linear, bisulcate
leaves, often to 3.5 cm long. Old individuals may become pendulous, with falcately ascending leaves.
Specimens examined:—Huila: Quebrada Ariari, above Galilea, 23 km ENE of Colombia, 2000 m, Fosberg
19617 (US).
Phlegmariurus polycarpos (Kunze) B.Øllgaard (2012b: 18).—Fig. 5A.
Lycopodium polycarpos Kunze (1835: 5).—Urostachys polycarpos (Kunze) Herter (1949: 76).—Huperzia polycarpos (Kunze) B.Øllgaard
(1987: 169). Type:—PERU: “Peruvia subandina, in sylv. apertioribus, Cassapi”, July 1829, Poeppig 1124 (holotype W).
Urostachys cuatrecasasii Herter (1954: 123).—Lycopodium cuatrecasasii (Herter) Morton (1964: 72). Type:—COLOMBIA. Valle
de Cauca: Cordillera Occidental, Vertiente Occidental, hoya del río Digua, Piedra de Moler, 900–1180 m, Cuatrecasas 15143A
(holotype US).
Illustrations: Øllgaard (1988: Fig. 19A); Lellinger (1989: Fig. 35).
Description:—Pendulous or recurved plants, with hanging divisions, to 30 cm long. Shoots homophyllous, almost
equally thick throughout, 10–15 mm in diameter including the leaves. Stems excluding leaves 1–1.5 mm thick near the
base, tapering to 0.5–1 mm, somewhat ridged by decurrent leaf bases, usually continuously sporangiate from 10–15
cm above the base and upward, to 6 times dichotomous. Leaves almost uniform throughout, densely crowded, borne
in alternating, irregular, often oblique whorls of 6–7 in basal divisions, upward of (3–)4, the whorls 0.5–1.5 mm apart,
forming (6–)8–14 indistinct longitudinal ranks, wide-spreading to ascending, often subsecund, usually with vertical
lamina due to a twist at the base, linear to filiform, 6–10(–12) × 0.4–0.5 mm, evenly tapering, usually oblique-falcately
curved, adaxially flat or shallowly canaliculate, often with an abaxially prominent vein near the leaf base, with green
decurrent leaf bases. Sporangia ca. 1 mm wide.
FIGURE 12. A. Phlegmariurus killipii (Herter) B. Øllg. (Schlim 1030, BR), growth habit. A2. Close-up of basal division, A2, terminal
fertile division.—B. Phlegmariurus pithyoides (Fosberg 19617, US), growth habit. B1. Closeup of leaf bases. B2. Diagrammatic cross
section of leaf. B3. Close-up of sporangiate division. (Del. K. Tind)

Distribution:—Costa Rica, Panamá, Colombia to Bolivia.
Habitat:—Epiphytic in cloud forest. Elev. 300–1200 m.
Notes:—Closely related to Phlegmariurus wilsonii, but usually pendulous, and smaller in all parts, with basally
twisted, falcately ascending leaves. From P. sarmentosus it is distinguished by the lack of leaf base auricles. From P.
mollicomus it differs in the more patent, twisted leaves, with much less prominent vein.
Specimens examined:—Valle del Cauca: Cisneros, Dagua Valley, 300–500 m, Killip 11472 (US). Río Timbiquí,
0–500 m, Lehmann 8908 (B, F, GH, K, US).
Phlegmariurus wilsonii (Underwood & F.Lloyd) B.Øllgaard (2012a: 481).—Fig. 13.
Lycopodium wilsonii Underwood & F.Lloyd (1906: 111).—Urostachys wilsonii (Underwood & F.Lloyd) Herter (1923: 163).—Huperzia
wilsonii (Underwood & F. Lloyd) B.Øllgaard (1987: 170). Type:—PUERTO RICO: Luquillo Mountains, Wilson 153 (lectotype NY,
here designated). Note: According to the protolog the type is Wilson 271, but Wilson 153 (NY) is annotated by the author as sp. nov.
and fits the description, while Wilson 271 (NY) is Phlegmariurus dichotomous and is not annotated as sp. nov.
Lycopodium andinum Herter (1909a: 49) non Rosenstock (1908: 239).—Lycopodium lindavianum Herter (1909b: 90).—Urostachys
lindavianus (Herter) Nessel (1939: 146).—Huperzia lindaviana (Herter) Holub (1985: 74). Type:—ECUADOR: “In territorio rei
publ. Ecuador”, Fraser s.n. (lectotype P [Herb. Drake], designated by Øllgaard 1988: 97; isotype, G).
Lycopodium trichodendron Herter (1909a: 49).—Urostachys trichodendron (Herter) Herter (1923: 163). Syntypes:—GUADELUPE:
L’Herminier s. n. (P00559253), Bory 103 (P00559254).
Lycopodium stamineum Maxon (1912: 2).—Urostachys stamineus (Maxon) Nessel (1939: 148). Types:—PANAMA. Chiriquí: above El
Boquete, ca. 1750 m, Maxon 5636 (holotype US).
Lycopodium arcanum Maxon (1938: 310). Type:—HONDURAS: Comayagua, above El Achiote , above plains of Siguatepeque, 1800 m,
Yuncker, Dawson & Youse 6149 (holotype US).
Urostachys mandiocanus (Raddi) Herter var. filifolius Herter (1954: 124). Type:—ECUADOR: Eastern slope of eastern cordillera, bank
of Palora river, 1500 m, Rimbach 125 (holotype US).
Illustrations: Øllgaard (1988: Fig. 1C); Mickel & Beitel (1988: Fig. 4B); Lellinger (1989: Fig. 44).
Description:—Plants erect, arcuate-spreading to pendulous, to 20(–30) cm long. Shoots homophyllous, 1.5–2.5(–3)

cm in diameter including leaves, equally thick throughout, or in some slender individuals gradually tapering to 1–1.5
cm in terminal divisions. Stems excluding leaves (1–)1.5–2(–3) mm thick at the base, often tapering to (0.7–)1 mm,
prominently ridged by decurrent leaf bases, pale green to stramineous, often with bright red spots on leaf bases,
sporangiate, often in seasonally produced zones, from 5–15 cm above the base and upward, 3–6(–9) times dichotomous.
Leaves usually uniform throughout, borne in alternating, often irregularly oblique whorls of 6–7, these 0.5–1.5 mm
apart in basal divisions, upward often in whorls of 4–5, forming 8–14 indistinct longitudinal ranks, perpendicularly
spreading to ascending, straight to upward curved, usually not twisted at the base, linear to filiform, (6–)10–17 ×
0.3–0.5 mm, quickly narrowed to ca. 0.2 mm wide due to involution, gradually tapering toward the tip, adaxially
canaliculate to involute, often with a prominent vein abaxially near the base. Leaves of terminal divisions in old plants
sometimes gradually reduced to 6(–4) mm long. Decurrent leaf bases usually not wider than the lamina base, often
bright red. Sporangia 1–1.5 mm wide.
Distribution:—Mexico (Oaxaca) to Panama, West Indies, northern South America south to Bolivia, Brazil (Mato
Grosso).
Habitat:—Epiphytic (rarely terrestrial) in montane forest, elev. 200–2700 m.
Notes:—Phlegmariurus wilsonii is related to P. dichotomus, P. mandiocanus (Raddi 1819: 280) B.Øllgaard
(2012a: 480) from Brazil, and P. pithyoides from Central America, Colombia, and Venezuela. With these it shares the
bottle-brush-like aspect, and with the latter two the bright red coloration of the leaf bases. Pendulous individuals of
P. wilsonii are rather similar to P. polycarpos, but the latter has shorter, more flattened, leaves, and uniformly falcate
leaves due to a basal twist, and this species apparently is always pendulous. Pendulous individuals of P. wilsonii may
be separated from P. sarmentosus by the lack of auricles at the lamina base.
The absence or presence of red colour on the leaf bases seems uncorrelated with other characters. Also the intensity
and size of the coloration is variable. The direction of leaves is correlated to growth habit. In erect-growing plants the
leaves are wide-spreading to slightly reflexed, while in pendent plants the leaves are somewhat ascending.
Several collections from Prov. Antioquia are unusually slender, but even larger plants among these are still erect-
growing and have longer leaves than in the closely related and spreadingly pendulous P. polycarpos.
Specimens examined:—Antioquia: Mun. Valdívia, Vereda San Fermín, 1 km along road Ventanas (Mun.
Yarumal) –Briceño, 1700 m, Callejas et al. 6201 (AAU). Juntas, Lehmann 5084 (K, US). Urrao, PN Las Orquideas,
Vereda Calles, Margen derecha del Río Calles, 1350–1450 m, Pipoly et al. 17749 (MO). Vereda Calles, PN Las
Orchideas, Margen derecha del Río Calles, en el filo NW de la cabaña de Calles, 1450 m, Cogollo et al. 7974 (MO).
Antadó, valley of Río San Jorge, PN Paramillo, 1560 m, Gentry & Cuadros 79014 (MO). Chocó: Macizo de Tamaná,
costado NE del río Tamaná, 2700 m, Torres et al. 2120 (COL). Río Salaqui, 6 days upstream Río Sucio, 200 m, Duke
11354 (NY). 5–8 kms E de Playa de Oro (E of Tadó), 140 m, Juncosa 2502 (AAU). Norte de Santander: Ocaña,
Lehmann s.n. (US). Cord. Oriental, Sarare region, drainage of Río Cubuzón, El Indio, 420–430 m, Cuatrecasas 13085
(F). El Valle: S. Antonio, 2000 m, 18 May 1939, Alston 8605 (BM).
FIGURE 13. Phlegmariurus wilsonii (Juncosa 2502, AAU). A. Growth habit. A1: Fertile terminal division.
Group of Phlegmariurus hartwegianus
It was a surprise to find the two distinctly heterophyllous species in this group to be associated with Phlegmariurus
hartwegianus, which was earlier assumed to be related to P. taxifolius. However, they are both obviously more robust
than the delicate species of the P. myrsinites group.
Phlegmariurus campianus (B.Øllg.) B.Øllgaard (2012b: 13). Fig. 14A.
Huperzia campiana B.Øllgaard (1988: 109). Type:—ECUADOR. Loja: E of Nudo de Cajanuma, Øllgaard et al. 57861 (holotype AAU;
isotypes LOJA, NY, QCA).
Illustration: Øllgaard (1988: Fig. 22B).
Description:—Rather robust, pendulous plants, at least to 1 m long. Shoots heterophyllous, the basal divisions, to 25–
60 cm from the base, ca. (15–)20–30 mm in diameter including the expanded leaves, then abruptly constricted to 2–4

mm thick including the imbricate, reduced leaves of the to 50 cm long, constricted, quadrangular terminal divisions.
Stems exclcluding leaves 2–4 mm thick at the base, tapering to 0.5–1.5 mm upward, pale greenish or brownish, at least
to 10 times dichotomous. Expanded leaves of basal divisions usually uniform throughout, borne in alternating whorls
of 3, these (2.5–)3–5 mm apart, forming 6 longitudinal ranks, spreading to perpendicular to the stem, lanceolate to
widely lanceolate-ovate, usually widest below the middle, (7.5–)10–15 × 2.5–4(–4.5) mm, often somewhat shorter
near the contraction of the shoot, twisted at the base, with flat to revolute margins, herbaceous to subcoriaceous, dull to
somewhat lustrous. Leaves of terminal constricted divisions decussate, or rarely in alternating whorls of 3 near the base
of constricted divisions, the pairs or whorls 1–2.5 mm apart, usually sporangiate throughout, appressed and clasping
with their bases, bluntly to sharply carinate, widely ovate to triangular-ovate, short-acuminate or mucronulate, 2–3 ×
1.5–2 mm, the sporophylls equalling or slightly exceeding the sporangia. Sporangia 1.2–2 mm wide.
Distribution:—Colombia to Peru.
Habitat:—Epiphytic in upper cloud forest and elfin forest, elev. 2400–3800 m.
FIGURE 14. Phlegmariurus campianus (Ecuador: Holm-Nielsen et al. 5339, AAU), growth habit. A1. Close-up of basal division. A2.
Close-up of terminal sporangiate division.—B. Phlegmariurus echinatus (Ecuador: Harling et al. 8462, GB) growth habit. B1. Close-up
of basal division. B2. Close-up of terminal sporangiate division (Ecuador: Camp E-5119A, AAU). Del. B. Johnsen.
Notes:—Phlegmariurus campianus resembles P. echinatus with respect to growth habit, but is generally smaller,
and especially the constricted terminal divisions are narrower and subterete, with leaves only 2–3 mm long.
Specimens examined:—Cundinamarca: La Calera , páramos altos, más al norte del llamado páramo de Palacio,
3500–3800 m, Uribe 5859 (COL). La Peña, Bogotá, Apollinaire 9 (US). Santander del Norte: Pamplona, 1911,
Aristé-Joseph s.n. (US)
Phlegmariurus echinatus (Spring) B.Øllgaard (2012b: 14).—Fig. 14B.
Lycopodium echinatum Spring (1849: 4).—Huperzia echinata (Spring) Trevisan (1874: 247).—Urostachys echinatus (Spring) Nessel
(1939: 178). Type:—COLOMBIA: “Mudo [?Muzo, Boyacá?], January 1826, Nova Granada”, Purdie s n. (lectotype K; isolectotypes
BM, GH, NY), designated by B.Øllgaard (1988: 109).
Lycopodium molongense Herter (1909a: 51).—Urostachys molongensis (Herter) Nessel (1939: 240).—Huperzia molongensis (Herter)
Holub (1985: 75).—Phlegmariurus molongensis (Herter) B.Øllgaard (2012b: 17). Type as to protologue: “Molong”, 1865, Pearce
s.n. (holotype K; isotype BM) [erroneously as “Ostaustralische Provinz: Gebirgswälder Neu-Süd-Wales”]. Pearce did not collect in
Australia, but extensively in the Andes. The type was probably collected in northern Peru.
Description:—Robust to very robust, pendulous, reported to 250 cm long. Shoots heterophyllous, usually not sharply
differentiated, 20–35 mm in diameter including the expanded leaves in the basal 1–30(–45) cm of the plant, upward
gradually to abruptly constricted to 3–8 mm in diameter including the shorter, imbricate leaves in the terminal divisions.
Stems excluding leaves 2–3.5 mm thick at the base, tapering to 1–1.5 mm, pale greenish or brownish, to at least 6 times
dichotomous. Expanded leaves of basal divisions borne in alternating whorls of 3, forming 6 longitudinal ranks, patent
to ascending or slightly recurved, lanceolate to lanceolate-ovate, widest near the base, acute, 10–18 × 3–6(–7) mm,
flat, or with slightly revolute margins, coriaceous, often lustrous, usually twisted at the base, the distance between leaf
whorls 3–6 mm. Leaves of constricted divisions decussate or subdecussate, imbricate, discontinuously sporangiate,
ovate to subcordate, acute, sharply carinate and clasping with the base, usually conduplicate in the apex, usually
lustrous, (2.5–)3–10 × 3–6 mm, the distance between leaf pairs 1–3 mm. Sporangia 1.5–2.5 mm wide.
Distribution:—Costa Rica, Andes from Venezuela to Northern Peru.
Habitat:—Montane, epiphytic in wet cloud forest, 2900–3500 m elev.
Notes:—This is one of the largest and most robust among the epiphytic species in the Andes. It is easily distinguished
because of its size, and the sharply quadrangular terminal divisions. The expanded leaves are sometimes developed
only along a short portion of the stem base, and are sometimes lacking in herbarium specimens, or disappeared in very
old individuals.
Phlegmariurus molongensis was earlier maintained as a species separate from P. echinatus (Øllgaard 1988, 1994)
because some of the ultimate branches in the type of the latter have 6-seriate, less sharply carinate leaves in the
gradually constricted terminal divisions. However, other ultimate branches of the type are sharply quadrangular and
decussate just as in material earlier referred to P. molongensis. I assume that the very large size of the type specimens
of P. echinatus may account for this difference of phyllotaxis.
The name Lycopodium meridense Mettenius was often used for Venezuelan material of this species (e.g. Nessel
1939), but the name is not found in “Novara Reise, Bot. 1” (1870), as cited by Herter (1949: 28), nor has it been validly
published in other references I have seen.
Specimens examined:—Cauca: Macizo Colombiano, páramo de las Papas, between Boquerón and La Hoyola,
3200–3510 m, Idrobo et al. 3575 (COL, G, NY). Macizo Colombiano, páramo de las Papas, around Laguna de
Cusiyaco, 3470 m, Barclay & Juajibioy 5962 (GH, MO). Cord. Central, W slope, headwaters of Río Palo, Río López
valley, Quebrada del Duende, 3400–3450 m, Cuatrecasas 18939 (GH). Cundinamarca: Road E of Guasca, 3200 m,
Haught 5896 (US). Mun. Fomeque, road to Páramo de Chingaza, 2900 m, Camargo & Huertas 7999 (COL). Nariño:
Pasto, Jameson 435 (BM, E, G, US). Cord. Oriental, Headwaters of Río Tellez, above Soledad, 3150 m, Ewan 16521
(GH, NO, P, S, UC, US). Alto de Tabano, Pasto, André 2962 (F, GH, NY, US). Páramo El Frailejonal, 8 km E of
Mocondino, 3400 m, Ramirez et al. 10428 (COL). Tolima: Road Mariquita—Manizales, “La Libia”, 3000–3200 m,
Santa & de Escobar 852, 853 (AAU, COL, HUA).

FIGURE 15. A. Phlegmariurus hartwegianus (Ecuador: Holm-Nielsen et al. 6362, AAU), growth habit. A1. Close-up of sporangiate
division.—B. Phlegmariurus myrtuosus (Colombia: Purdie s.n. K holotype), growth habit. B1. Close-up of sporangiate division.—C.
Phlegmariurus rosenstockianus (Ecuador: Camp 5187, AAU), growth habit of terrestrial plant. C1. Close-up of sporangiate division—D.
Phlegmariurus rosenstockianus (Ecuador: Wiggins 10324, US), growth habit of pendulous plant. D1. Close-up of basal division. D2.
Close-up of sporangiate division. Del. B. Johnsen
Phlegmariurus hartwegianus (Spring) B.Øllgaard (2012b: 15).—Fig. 15A.
Lycopodium hartwegianum Spring (1849: 14).—Huperzia hartwegiana (Spring) Trevisan (1874: 248).—Urostachys hartwegianus
(Spring) Nessel (1939: 90). Type:—COLOMBIA: Prov. Popoyan, Hartweg 1466 (lectotype K with annotation by Spring, designated
by Øllgaard (1988: 76); isolectotypes BM, G, LG, NY).
Lycopodium funckii Herter (1909a: 44).—Urostachys funckii (Herter) Nessel (1939: 70). Type:—VENEZUELA. Mérida: Culata, 8000
pds. [ca. 2438 m], Funck & Schlim 1568 (holotype P00559195; isotypes BM, BR).
Lycopodium caracasicum Herter (1909b: 88).—Urostachys caracasicus (Herter) Nessel (1939: 90).—Huperzia caracasica (Herter) Holub
(1985: 71). Type:—VENEZUELA: Caracas, Jan. 1856 (fide protol.), Gollmer s.n. (holotype B).
Urostachys maxonii Nessel (1940a: 165). 1940. Type:—ECUADOR. Imbabura: Otavalo, Spruce 10 125 (holotype P-00559192; isotype
BONN [fragment, Herb. Nessel 422]).
Illustrations: Øllgaard (1988: Fig. 14A).
Description:—Robust, recurved or pendulous plants, at least to 90 cm long. Shoots homophyllous to gradually

heterophyllous, 15–20 mm in diameter including the leaves in basal divisions, tapering to (6–)8–12 mm in diameter
in terminal sporangiate divisions, sometimes not or slightly tapering (juvenile and tardily sporangiate plants). Stems
excluding leaves 2.5–3.5(–5) mm thick at the base, tapering to 1.5–2 mm thick upward, pale greenish or brownish, to
6(–9) times dichotomous, sporangiate from 20–30 cm above the base and upward. Leaves of basal divisions borne in
alternating whorls of 4(–5), these 2–4 mm apart, forming 8(–10) usually distinct longitudinal ranks, falcately patent-
ascending to imbricate, radially directed to somewhat secund, upward almost completely covering the stem and
sporangia, lanceolate, firmly herbaceous to rigidly coriaceous, usually lustrous, 14–20 × 2.5–3.5 mm, widest below
the middle to near the base, convex at the base beneath, usually slightly concave toward the tip, with smooth, flat to
somewhat involute margins, the vein slightly prominent at the base and apically obscure. Leaves of terminal divisions
conform, or usually smaller, 7–10(–13) × 1.5–3 mm, often somewhat clasping with the widened base, softer and less
lustrous. Sporangia 1.7–3 mm wide.
Distribution:—Hispaniola, southern Mexico, Guatemala, Andes from Venezuela to Peru. Ramírez Padilla &
Macias Pinto (2007) reported this species from Dept. Cauca.
Habitat:—Epiphytic or rupestral in montane forest, or in exposed habitats, 1500–3800 m.
Notes:—Phlegmariurus hartwegianus resembles P. taxifolius and earlier was placed in the same group. However,
molecular data (Testo et al. 2018) suggest a closer relation to P. echinatus and P. campianus, two distinctly heterophyllous
species.
Phlegmariurus hartwegianus is a widely distributed and somewhat variable species. The variation seems partly
correlated with habitat type. It often occurs in habitats that are too dry for the other epiphytic species. In relatively dry
and exposed rupestral sites, the plants initially grow erect, then recurved, and may develop rather short and rigid shoots
only. Such forms were recognized as Lycopodium caracasicum by Herter and others. In moist, shaded, epiphytic
habitats long, lax, pendulous forms are developed.
This is a robust species, usually with strongly coriaceous, nitid leaves in alternating whorls of four. In Colombia
it varies from a long, lax, epiphytic growth habit to initially erect and recurved in lava rocks.
Specimens examined:—Boyacá: Mun. Villa de Leiva, Vereda La Capilla, rocky bank of Río Iguaque, 6 km from
La Capilla, 2200 m, Stancik et al. 980 (AAU). César: Sierra de Perijá, 6 km E of Manaure, 42 km E of Valledupar,
7 km from Venezuelan Border, 2175 m, Grant 10753 (US). Sierra Nevada de Santa Marta, SE slopes, drainage of
Río Donachui, Meollaca, 3450–3550 m, Cuatrecasas & Castañeda 24650 (COL, US). Cundinamarca: Bogotá near
Cuervo, Triana s. n. (BM). Bogotá, Amórtegui A56 (US). Guajira: Riohacha (Sierra Nevada), 3050 m, Schlim 878
(BR, G, P). Páramos of Sierra Nevada de Santa Marta, ca. 30 mi inland from Dibulla, 3840 m, Seifriz 447 (US).
Magdalena: Sierra Nevada de Santa Marta, Cerro Quemado trail, 1500–2300 m, Martin 3690 (US). Santander: Near
La Baja, 2700 m, Killip & Smith 18329 (GH, US).
Group of Phlegmariurus lindenii
Phlegmariurus lindenii (Spring) B.Øllgaard (2012b: 16).—Fig. 1C.

Lycopodium lindenii Spring (1849: 27).—Urostachys lindenii (Spring) Nessel (1939: 177).—Huperzia lindenii (Spring) Trevisan (1874:
247). Type:—COLOMBIA. Tolima: Prov. Mariquita, Páramo du Tolima, elev. 12–13000 ped., Linden 1002 (lectotype LG, isotype
BR, P) designated by Øllgaard (1988: 84).
Lycopodium stuebelii Herter (1909a: 53).—Urostachys stuebelii (Herter) Nessel (1939: 241).—Huperzia stuebelii (Herter) Holub (1985:
77). Type:—COLOMBIA. Nariño: Volcán de Cumbal, oberhalb Cumbal, Stübel 441 (holotype B).
Lycopodium sodiroanum Herter (1909b: 91).—Urostachys sodiroanus (Herter) Nessel (1939: 240).—Huperzia sodiroana (Herter) Holub
(1985: 77). Type:—ECUADOR. Tungurahua: E slopes of Volcán Tungurahua, 3500 m, Rimbach 28 (P00559131, S, US).
Urostachys wohlberedtii Nessel (1935: 69).—Lycopodium wohlberedtii (Nessel) Morton (1964: 73).—Huperzia wohlberedtii (Nessel)
Holub (1985: 78). Type:—ECUADOR. Pichincha: Malchingui, 3000 m, Hitchcock s. n. (lectotype herb. Nessel, BONN) designated
by Øllgaard (1988: 84).
Description:—Plants flaccidly pendulous, at least to 70 cm long. Shoots usually gradually heterophyllous, often
homophyllous, 10–18 mm in diameter including leaves in basal divisions, tapering to 5–10 mm in terminal divisions.
Stems excluding leaves 1–1.5(–2) mm thick at the base, tapering to 0.5–0.8 mm upward, pale greenish to brownish,
sporangiate from 15–30 cm above the base and upward, often more than 10 times dichotomous. Leaves of basal
divisions borne in alternating whorls of 3–4, these (1–)2.5–5 mm apart, gradually more crowded toward the apex,
patent-ascending, sometimes slightly secund, linear-lanceolate, widest in or below the middle, short-acute, abaxially
flat to slightly convex, 8–11 × 1.3–2 mm, firmly herbaceous to subcoriaceous, dull to slightly shining, with obscure to
prominent vein, with smooth margins. Leaves of terminal divisions sometimes conform, but usually shorter and wider,
borne in whorls of 3, lanceolate to ovate or subdeltoid, somewhat clasping with the widened base, suddenly tapering
into a short to long acumen, acute, (2.5–)4–7 × 1.3–2.5 mm, abaxially convex. Sporangia 1.5–2 mm wide.
Distribution:—Colombia and Ecuador.
Habitat:—Epiphytic and common in forest near the timber line, or in moss cushions on rocks in páramo, and
often on solitary trees in the páramo. Elev. 3400–4300 m.
Notes:—The number of heterotypic synonyms partly reflect the variability of the species in response to
environmental differences. Molecular data (Testo et al. 2018) indicate an isolated position of this species in the group
including Phlegmariurus taxifolius.
Specimens examined:—Antioquia: Sonson, 1871, Wallis s.n. (HBG). Boyacá: Páramo de la Rusia, NWN of
Duitama, Serrania Peña Negra, 3970 m, Cleef 7431 (COL). Nariño: Ascent to Cumbal, 3400–4300 m, Hagemann &
Leist 1977 (COL). Putumayo: Banks of Putumayo at Concepción, Rimbach 229 (US). Risaralda: Mun. Sta Rosa de
Cabal, Termales de Sta Rosa–Hda. La Sierra, 3670 m, Wolf 848 (COL). Parque Los Nevados, near Finca La Sierra,
3650 m, Verweij & Beukema F 30hh (U).
Group of Phlegmariurus taxifolius
Phlegmariurus taxifolius (Swartz) Löve & Löve (1977: 324).—Fig. 16.
Lycopodium taxifolium Swartz (1788: 138).—Huperzia taxifolia (Swartz) Trevisan (1874: 248).—Urostachys taxifolius (Swartz) Herter
(1923: 162). Type:—A remounted specimen, marked as type, but without original annotation, in the type herbarium in S, is possibly
the holotype.
Lycopodium passerinoides Kunth (1816: 41).—Huperzia passerinoides (Kunth) Trevisan (1874: 248).—Urostachys passerinoides (Kunth)
Nessel (1927: 417).—Urostachys rubiginosus Nessel (1940a: 162). Type:—PERU. Jaén: “prope Olleras et Aipate”, elev. 747 hexp.,
Humboldt s.n. (holotype P-Humb. P00669228, isotypes P-00559263, BONN-herb. Nessel 160).
Lycopodium nitens Schlechtendal & Chamisso (1830: 623).—Huperzia passerinoides (Kunth) Trevisan var. nitens (Schlecht. & Cham.)
Trevisan (1874: 248).—Urostachys nitens (Schlecht. & Cham.) Herter (1923: 164).—Urostachys passerinoides (Kunth) Nessel var.
nitens (Schlecht. & Cham.) Nessel (1927: 418).—Urostachys schwendeneri (Herter) Herter var. nitens (Schlecht. & Cham.) Nessel
(1939: 166). Type:—MEXICO. Jalapa: ”In arboris vetustis prope Jalapam”, Schiede & Deppe s. n. (holotype B, isotype BM).
Lycopodium herminieri Spring (1841: 514).—Urostachys herminieri (Spring) Herter (1923: 162). Type:—GUADELUPE: L’Herminier s.
n. (holotype P00559265).
Lycopodium schwendeneri Herter (1909a: 50).—Urostachys schwendeneri (Herter) Herter (1923: 165). Type:—MEXICO: (indicated as
locus classicus by Herter 1923). No type designate
Urostachys bruelkei Nessel (1934: 184). Type:—COLOMBIA: Colombia, 1600—2100 m, E. P. Killip 11595 (holotype BONN herb.
Nessel 399).
Lycopodium cubanum Herter (1909a: 50).—Urostachys cubanus (Herter) Herter (1923: 165).—Huperzia cubana (Herter) Holub (1985:
71). Type:—CUBA: “El Gato, St Yago de Cuba, 1844”, Linden 2185 (P! 2 sheets, both annotated later by Herter).
Illustrations: Lellinger, 1989: Fig. 39; Mickel & Beitel, 1988: Fig. 4A; Øllgaard, 1988: Fig. 15A
FIGURE 16. Phlegmariurus taxifolius A: (Colombia: Espinal 3170, COL), growth habit. A1. Close-up of sporangiate division—B.
Phlegmariurus taxifolius (Espinal 3109, COL) growth habit. B1. Close-up of sporangiate division.
Description:—Lax and pendulous, or sometimes recurved from an erect and somewhat rigid stem base, up to 50–70
cm long. Shoots usually gradually tapering from ca. 20–30 mm in diameter incl. leaves at the base, to (3–)4–7 mm
in diam. in terminal, densely sporangiate divisions of fully developed plants, sporangiate from 10–50 cm above the
base and upward, sometimes not, or only slightly tapering (juvenile or tardily sporangiate individuals), rarely abruptly
constricted. Stems excluding leaves 1.5–3 mm thick at the base, tapering to 1–1.5 mm upward, somewhat ridged by
decurrent leaf bases, pale greenish to brownish, usually 4–5(–7) times dichotomous. Leaves usually reduced and
modified upward, borne in alternating whorls or irregular low spirals of 3 or 4, these 1.5–4 mm apart, forming 6–8

indistinct longitudinal ranks. Leaves of basal divisions spreading to ascending or somewhat appressed, often twisting
the lamina to a vertical position from the leaf base, narrowly lanceolate, broadest in the lower half, broadly joined to
the stem, firmly herbaceous to subcoriaceous, 14–23 × (1.5–)2–3.5 mm, almost flat or somewhat concave adaxially,
with flat or slightly revolute, smooth margins, with evident to somewhat prominent vein abaxially. Leaves of middle
and terminal divisions usually gradually shorter, narrower and more appressed, abaxially more convex, with involute
margins. Leaves of fully sporangiate divisions often distinctly 6-ranked, with strongly broadened, clasping base, partly
covering the sporangia, often abruptly contracted into a short to long, narrow, involute apex, 3–8 mm long, 1–1.5(–2)
mm wide. Sporangia 1–1.5 mm wide.
Distribution:—Central America, West Indies, northern South America, south to Bolivia and Brazil (Paraná).
Habitat:—Epiphytic in lower to mid-elevation wet montane forest, elev. 100–3000 m.
Notes:—As testified by the number of synonyms Phlegmariurus taxifolius is a variable species. Intermediates
between P. taxifolius, P. linifolius, and P. homocarpus are frequent.
Specimens examined:—Caldas: Salento, 3000 m, Sneidern 3231 (S). Cauca: Cajibío, 1850 m, Espinal 3170
(COL), 3169 (COL). Quebrada La Pita, SW of Tambo, 1100 m, Haught 5271 (US), 5272 (COL, US). César: 6
kms E de Manaure, 2100 m, Grant 10749 (US). Sierra de Perijá, E of Manaure, Hda Nuevo Horizonte, El Podrido,
1550—1600 m, Cuatrecasas & Romero Castañeda 25390 (COL, US). Chocó: Road Río Macana—Alto de Macana,
100—500 m, Gentry & Juncosa 41034 (AAU, COL, MO). Cundinamarca: Usaquén, March 1905, Frére Apollinaire
s.n. (E, K). Guajira: Santa Marta, Horqueta Mts., 1980 m, Smith 2601 (GH, NY). Rio Hacha, Sierra Nevada, 3050
m, Schlim 879 (BR, F, G). Magdalena: Around San Andrés de la Sierra, W slope of Cord. de Santa Marta, 1100–1300
m, Pittier 1702 (US). Santa Marta, Sierra de Onaca, 1525 m, Smith 2471 (NY). Sierra Nevada de Santa Marta, road
to crest of San Lorenzo, 1700 m, Santa & Escobar 651 (AAU). Sierra de Perijá, 6 km E-NE of Manaure, 42 km E of
Valledupar, 7 km from Venezuelan border, 2100 m, Grant 10749 (US). Sierra de Perijá, E of Manaure, Hda. Nuevo
Horizonte, El Podrido, 1550–1600 m, Cuatrecasas & Romero Castañeda 25390 (COL). Norte de Santander: Ocaña,
Schlim 469 (P). Ocaña, 2440 m, 1846—1852, Schlim 1031 (BR). Ocaña, 3350 m, Schlim 1028 (BR, F, G, K). Ocaña,
3050 m, Schlim 643 (BR, G, K, P). Risaralda: Mun. Santa Rosa de Cabal, W of Tremales de Santa Rosa, Hda La
Gaviota, 1980 m, Wolf 1092 (COL, U). Province unknown: 1871, Karsten s.n. (HBG); Kalbreyer 463 (K).
Group of Phlegmariurus linifolius
Phlegmariurus homocarpus (Herter) B.Øllgaard (2012b: 15).—Fig. 17A.
Urostachys homocarpus Herter (1923: 163).—Lycopodium homocarpum (Herter) C. Christensen (1937: 92).—Huperzia homocarpa
(Herter) Holub (1985: 73). Type:—DOMINICAN REPUBLIC: La Vega: “In sylvis humidis”, 1300 m, Padre Miguel Fuertes 1785
(holotype B; isotypes BONN-Nessel 354, GH, NY, P).
Description:—Plants flaccidly pendulous, at least to 150 cm long. Shoots homophyllous, or the leaves slightly
modified toward the shoot apices, 15–20 mm in diameter including leaves throughout, or tapering to 7–10 mm in
apical divisions. Leaves borne in alternating, irregular whorls of 3–4, these 2–5 mm apart, forming 6–8 indistinct
longitudinal ranks, sometimes densely crowded in sporangiate divisions, spreading to ascending, sometimes loosely
appressed in apical divisions, linear to linear-lanceolate, in apical divisions widened just above the base to near the
middle, from there evenly tapering into a long narrow tip, adaxially convex, abaxially concave with slightly prominent
vein, 10–14 mm long in basal divisions, sometimes reduced to (7–)8–10 mm long in apical divisions, 1–1.5 mm wide,
softly to firmly herbaceous, dull to somewhat shining, with slightly revolute, smooth margins, usually twisted at base
and with slightly falcate-ascending, vertical lamina. Sporangia 1.5–2.2 mm wide.
Distribution:—Total distribution uncertain, but including Greater Antilles, Costa Rica, Panama, and northern
Andes.
Habitat:—Epiphytic in upper montane forest, elev. 2800–2900 m, but probably with a wider range of elevation.
Notes:—Homophyllous, lax, pendulous, with rather distant whorls of relatively short, linear-lanceolate, usually
basally twisted leaves. Difficult to delimit in relation to Phlegmariurus taxifolius, P. linifolius, and P. dichotomus.
Specimens examined:—Cundinamarca: Near Bogotá, Mangonos , 2900 m, Lindig 1520 (K, P). Nariño:
Drainage of upper Río Tescual, above Córdoba, 2840–2890 m, Ewan 16296 (BM, COL, NO, UC, US).
FIGURE 17. A. Phlegmariurus homocarpus (Ecuador: Prieto P-224, AAU) growth habit. A1. Close-up of sporangiate division.—B.
Phlegmariurus linifolius var. linifolius (Ecuador: Holm-Nielsen et al. 2734, AAU) growth habit. B1. Leaf of basal division. B2. close-up
of middle sporangiate division. B3. Close-up of terminal sporangiate division.—C. Phlegmariurus linifolius var tenuifolius (Holm-Nielsen
et al. 4422, Ecuador, AAU), growth habit. C1. Close-up of basal division. C2. close-up of sporangiate division. Del B. Johnsen
Phlegmariurus killipii (Herter) B.Øllgaard (2012a: 480).—Fig. 12A.
Urostachys killipii Herter (1954: 128).—Lycopodium killipii (Herter) C. Morton (1964: 72).—Huperzia killipii (Herter) B.Øllgaard (1987:
169). Type:—COLOMBIA: El Valle, La Cumbre, 1600–2100 m, Killip 11587 (holotype US 1189919; isotype GH).
Description:—Plants flaccidly pendulous, slender, at least to 50 cm long, at least to 10 times dichotomous. Shoots
nearly homophyllous to gradually heterophyllous, tapering from ca. 10–18 mm in diameter including leaves at the

base, to 3–5(–7) mm in terminal divisions, sometimes only slightly tapering. Stems excluding leaves 1–1.5 mm thick
at the base, tapering to 0.5–0.8 mm upward. Leaves uniform throughout or gradually modified, borne in irregular,
alternating whorls of 3 (–4), these 1–3 mm apart, in terminal divisions sometimes subdecussate. Leaves of basal
divisions spreading to ascending or loosely appressed, sometimes twisted at the base, linear-oblanceolate to elliptic,
narrowly cuneate, acute to short-acute, 8–11(–12) × (1–)1.2–1.8(–2.5) mm, almost flat or with slightly revolute
margins, softly to firmly herbaceous, with smooth margins. Leaves of distal divisions conform or gradually shorter,
narrower and more appressed, or abaxially convex, with involute margins. Leaves of sporangiate divisions 6-ranked or
subdecussate, rarely with a clasping base and contracted into a short, involute, bluntly carinate apex, 2–5 × ca. 0.6–1.5
mm. Sporangia ca. 1.5 mm wide.
Distribution:—Dominican Republic, Venezuela, Colombia, Ecuador, French Guyana.
Habitat:—Pendulous epiphyte in mid-elevation montane forest, elev. 1600–3000 m.
Notes:—A delicate species resembling P. tubulosus Maxon (Central America), but often completely homophyllous,
with elliptic to oblanceolate, usually to 10 mm long proximal leaves. Specimens examined:—Cauca: La Capilla, 25
km N of Popayán, 1740 m, Killip 38465 (US). Cundinamarca: Road Subatá–San Francisco, 2000 m, Haught 6105
(S, US). Huila: Mun. La Plata, Agua Bonita, finca Meremberg, forest of “Ojo de agua”, 2480 m, Torres 1183 (COL).
Norte de Santander: Ocaña, 2440–3050 m, Schlim 1030 (BR, G, K, P). Valle del Cauca: La Cumbre, 1600–2100 m,
Killip 11587 (GH, US).
Phlegmariurus linifolius (Linnaeus) B.Øllgaard (2012a: 480).—Figs. 17–19.
Lycopodium linifolium Linnaeus (1753: 1100).—Huperzia linifolia (Linnaeus) Trevisan (1874: 248).—Urostachys linifolius (Linnaeus)
Herter (1923: 165). Lectotype:—Martinique or Hispaniola, Plumier, Traité Fougères (1705: t. 166, Fig. C) designated by Proctor
(1977: 26).
Illustrations: Mickel & Beitel (1988: Fig. 4C); Øllgaard (1988: figs. 16B); Lellinger (1989: Fig. 30).
Description:—Plants pendulous, usually with flaccidly hanging divisions, the terminal divisions often aggregated in
fasciculate clusters, to 60(–80) cm long. Shoots homophyllous or gradually heterophyllous, equally thick throughout,
20–30(–45) mm in diameter including leaves, or gradually tapering to (7–)10–15 mm in diameter in terminal, densely
sporangiate divisions. Stems excluding leaves 0.5–1 mm thick at the base, slightly tapering upward, almost straight
to somewhat flexuous, pale greenish in life, discontinuously sporangiate from 10–30 cm above the base and upward,
to 5(–8) times dichotomous. Leaves of basal divisions spirally arranged, single, or in occasional pairs or whorls of 3,
not predominantly whorled, forming ca. 6 indistinct longitudinal ranks, subdistant, 1–5 mm apart, soft-herbaceous,
wide-spreading to ascending, straight to slightly falcate, usually with the lamina vertical due to a twist of the lamina
base, linear-lanceolate to broadly lanceolate, widest in the basal third or fourth, distinctly narrowed into a petiole-like,
twisted, usually perpendicular to deflexed lamina base, (10–)13–24 × 1–5 mm, flat, or with slightly revolute, smooth
margins. Leaves of middle and terminal divisions spirally arranged, paired or borne in irregular to regular, alternating
whorls of 3, conform, or usually narrower, (5–)10–15(–20) × (1–)1.5–2(–3) mm, often with a long narrow apex from
a subauriculate, non-twisted lamina base. Sporangia 1–1.5 mm wide.
Distribution:—Southern Mexico to Panama, West Indies, northern South America, south to Bolivia and
Amazonian Brazil.
Notes:—Phlegmariurus linifolius is a widespread and polymorphic species. Its variation is especially complex in
the northern Andes. The varieties keyed out below are fairly well characterized morphologically and are geographically
well defined. However, material from mid-altitudes, especially from the Andean slopes in Ecuador and Colombia,
discussed under P. linifolius var. tenuifolius, is difficult to define, including a series of intermediates between P.
linifolius, P. taxifolius, and P. homocarpus.
Four varieties are recognized: The type variety occurs in Central America, the West Indies, and along the Pacific
slopes of the Andes south to Ecuador. Variety tenuifolius (Nessel) B.Øllgaard occurs mainly along the lower eastern
slopes of the Andes, var. planifolius (B.Øllgaard) B.Øllgaard in Venezuela to Ecuador, while variety jenmanii (Underw.
& Lloyd) B.Øllgaard occurs in the Amazonian lowlands, in the Guyanas and along the northeastern Atlantic coast of
Brazil.
Key to the varieties of Phlegmariurus linifolius
1. Widest leaves of basal divisions lanceolate to widely lanceolate, (2–)2.5–5 mm wide .....................................................................2
–. Widest leaves of basal divisions linear-lanceolate, less than 2 mm wide ..........................................................................................3
2. Leaves of basal divisions uniformly twisted to a vertical position, slightly falcately curved and slightly acuminate.........................
.................................................................................................................................................................................... A var. linifolius
–. Leaves of basal divisions transverse, not twisted, generally straight and symmetrical, evenly narrowing to the apex .......................
................................................................................................................................................................................ D. var. planifolius
3. Shortest sporophylls of densely sporangiate divisions with distinctly widened, usually not twisted lamina base, usually ascending
to loosely appressed, matte green. Stem usually opaque-stramineous ................................................................... B. var. tenuifolius
–. Shortest sporophylls of densely sporangiate divisions usually conform to leaves of basal divisions, or shorter, usually with a
twisted, narrow to slightly widened lamina base, perpendicular to spreading-ascending, pale, transparently green or brownish
green. Stem, at least in terminal divisions, usually transparently stramineous to reddish brown, the vascular tissue usually visible
through the cortex ...................................................................................................................................................... C. var. jenmanii
A. Phlegmariurus linifolius var. linifolius—Fig. 17B.
Illustrations: Øllgaard (1988: Fig. 16B).
Habitat:—Epiphytic in lowland, lower montane rain forests, and cloud forest, from sea level to ca. 0–2000 m. Mainly
on the lower Pacific slopes and foothills of the Andes, common on Citrus trees.
Specimens examined:—Antioquia: Mun. Zaragoza, road to Segovia, 200 m, Santa 56l (AAU, COL, HUA,
MEDEL). Mun. Anorí, Corr. Providencia, Tirana Creek, at confluence with Río Anorí, 500 m, Soejarto et al. 3179
(COL). Mun. Mutatá, left bank of Río Mutatá, 180–200 m, Fonnegra et al. 2082 (AAU). Near Antioquia, Las Juntas,
1220 m, Wallis s.n. (HBG). Chigorodó, road to Turbo, 100–200 m, García Barriga 17632 (COL, NY, US). Cauca:
Munchique, 2100 m, Alston 8201 (BM). Pacific coast, Río Micay, Guayabal, 5–20 m, Cuatrecasas 14159 (US). Chocó:
Río Atrato, Yuto, rocky margins of river above Yuto, 70 m, Cuatrecasas & Llano 24136 (BR, K, S, SI, UC, US). Río
San Juan, between Palestina and Agua Negra, 5 m, Cuatrecasas 21554 (F, US). Corcovada Region, Upper Río San
Juan, ridge along Yeracuí Valley, 200–275 m, Killip 35346 (US). Negría, ca. 50 m, along Río San Juan, Killip 35020
(US). Lower Río Truandó, 3–12 km SSW of Río Sucio, 2–4 m, Lellinger & de la Sota 604 (US). Quibdó, banks of
río Ichó, Santa et al. 1156 (AAU, COL, HUA, MEDEL). Las Palisadas, on the Truandó, Schott s.n. (F). Near Quibdó,
81 m, Scolnik 1626 (US). Río Atrato below Quibdó, 55 m, Mägdefrau 1491 (COL). Cirurquita(?), 2000 m, Triana
s.n. (US1593677). Chocó: Cienegueta (?), 2000 m, Triana 696 (COL). Cundinamarca: Sasaima?, Lasaima?, 1200
m, Huertas & Camargo 510 (F). Guajira: Sierra Nevada de Santa Marta, between Río Hacha and Pueblo Viejo, 200
m, Barclay & Juajibioy 6841 (MO, US). Magdalena: Santa Marta, Doce Amo, Smith 2240 (NY). Nariño: Ricaurte,
La Planada, 1900 m, Benavides 699 (MO). Barbacoas, Lehmann 4078 (G). Mun. Espriella, Road to Tumaco before
Guayacana, 450 m, León & Maas 1513 (U). Santander: Near Barranca Bermeja, Magdalena Valley between Sogamoso
and Colorado Rivers, 100–500 m, Haught 1501 (COL, GH, UC, US). Valle del Cauca: Pavas, 1600–2000 m, Killip
11675 (US) Finca La Pradera, ca. 6 km SW of El Cairo on trail to Río Blanco, El Brillante–Boquerón, 2150–2200 m,
Lellinger & de la Sota 795 (US). Mun. Dagua, old road Buenaventura–Cali, Río Digua, quebrada Piedras de Moler,
610 m, Santa & D. Sánchez 1059 (AAU). Río Cali, El Silencio, 2000 m, Duque-Jaramillo 489 (COL). Buenaventura,
corr. Córdoba, banks of Río Dagua, 100 m, Santa & Hoyos 748 (AAU). Pacific coast, Río Cajambre, Barco, 5–80 m,
Cuatrecasas 16979 (F, US). Pacific coast, Río Yurumanguí, El Aguacate, 5–10 m, Cuatrecasas 16148 (COL, F, NY,
UC, US). Buenaventura, Bajo Calima, Río Calima, 40 m, Santa & Sánchez 1049 (AAU, COL, HUA). Buenaventura,
Hagemann 488 (COL). Río Cajimbe, near sea level, Haught 5355 (COL). Valle del Cauca/Quindio: Buenavista–
Quindió, André 2133 (K). Province unknown: Mutis 5365 (G). Kalbreyer 354 (K).
B. Phlegmariurus linifolius (Linnaeus) B.Øllgaard var. tenuifolius (Nessel) B. Øllgaard (2012b: 16).—Fig. 17C.
Urostachys linifolius (Linnaeus) Herter var. tenuifolius Nessel (1940a: 164).—Huperzia linifolia (Linnaeus) Trevisan var. tenuifolia
(Nessel) B.Øllgaard (1988: 90). Type:—PERU. Huánuco: Pampayacu, Poeppig [»Moritz 210 und 1155«] (lectotype BONN [Herb.
Nessel 381], designated by Øllgaard (1988: 90); isotype G).
Distribution:—Colombia to Bolivia.
Habitat:—Wet forests mainly of the Eastern Andean slopes, elev. 1000–2700 m.

Notes:—In the original publication of this variety, the protologue indicates collections by Funck & Schlim,
and by Moritz (Herb. Greven) from Venezuela, but does not cite collection numbers or locality data. These are in
the reference in Bärlappgewächse p. 162: “Venezuela: Funck & Schlim n. 1114—n. 210 und n. 1155 Moritz (Herb.
Greven)—Guiana: in feuchten Wäldern (Schomburgk)”. Sheet no. 381 in Herb. Nessel (BONN) includes 4 collections,
all annotated “Urostachys linifolius var. tenuifolius (Schomb.)” by Nessel. Among these are Funck & Schlim 1114
and “Moritz 210 und 1155”. The latter specimen is illustrated in Bärlappgewächse as cited above, and is chosen as
lectotype. The label of the lectotype is obviously wrongly copied from other material. A duplicate of this collection
is in Genève (G), and bears the label text: “no. 210 Lycopodium linifolium L. var. tenuifol. 1135”.” Perou Poeppig”
has been added in a different handwriting. The two duplicates are quite identical, including the epiphytic Hepatics
and Lichens found on them. Thus the label discrepancies may serve to illustrate the degree of accuracy one should be
prepared for in Nessel’s work. According to Kunze (1834: 4), the origin of this collection is Peru: montane forests at
Pampayacu (prov. Huánuco).
Specimens examined:—Cauca: Open forests around Popoyan, 1600–2200 m, Lehmann K.K. 20 (F, K, UPS).
Near Popoyán, 1740 m, Hno Azael s.n. (F, MO). Ca 8 km E of Popayán, 2000 m, Churchill & Rengifo 17316 (AAU).
Santander: Pamplona, Río Fuio, Kalbreyer 761 (K). Valle del Cauca: Cord. Occidental., W slope, Río Digua
drainage, Piedra de Moler, 900–1100 m, Cuatrecasas 15055 (F, US). Hoya del río Cali, Pichindé, Alto de las Brisas,
2160 m, Cuatrecasas 18282 (S). E slope, hoya del Río Cali, at union of ríos Pichindecito and Pichindé, 1580–1650 m,
Cuatrecasas 18761 (F, US)
C. Phlegmariurus linifolius (Linnaeus) B.Øllgaard var. jenmanii (Underwood & Lloyd) B.Øllgaard (2012a: 480).—
Fig. 18A.
Lycopodium jenmanii Underwood & F. Lloyd (1906: 112).—Urostachys jenmanii (Underwood & F. Lloyd) Nessel (1939: 158).—Huperzia
jenmanii (Underwood & F. Lloyd) Holub (1985: 74).—Huperzia linifolia (Linnaeus) Trevisan var. jenmanii (Underwood & F.
Lloyd) B.Øllgaard & P. G. Windisch (1987: 13). Type:—BRITISH GUYANA: Moruca River, Jenman s. n. (holotype NY, isotypes
BONN [Herb. Nessel 388], E).
Distribution:—Throughout the Amazonian lowland region of Venezuela, Colombia, Ecuador, Peru and Bolivia, the
Guayanas and Northern coastal Brazil south to São Paulo.
Habitat:—Lowland Amazonian forest to 600 m elevation.
Specimens examined:—Amazonas: Trapecio Amazonico, Atacuari River, 100 m, Schultes & Black 8568 (GH).
Norte de Santander: Región of Sarare, drainage of río Cubugón, El Indio, 420–480 m, Cuatrecasas 13085 (COL,
US). Vaupés: Río Kubiyu (Cubiyu), 200 m, 1970, Soejarto & Lockwood 2398 (COL, F, K). Río Negro near Piedra de
Cocuí, Caatinga, Schultes & López 9514 (US). Vichada: Comisaría Vichada, Río Vichada near San José de Ocune, 100
m, Haught 2808 (US).
D. Phlegmariurus linifolius (Linnaeus) B.Øllgaard var. planifolius (B.Øllg.) B.Øllgaard (2012b: 16).—Fig. 19.
Huperzia linifolia (Linnaeus) Trevisan var. planifolia B.Øllgaard (2004: 35–37). Type:—COLOMBIA. Magdalena: Sierra Nevada de
Santa Marta, forest between Finca Risaralda and Finca Los Arroyitos, 73°58’W 10°56’N, 1700–1900 m, Kirkbride, 2293 (holotype
COL; isotypes COL, NY, US).
Illustrations:—Øllgaard (2004: Fig. 4).
Distribution:—Andes of central Ecuador to Colombia and Venezuela, in Sierra Nevada de Santa Marta, and the Sierra
de Perijá in northernmost Colombia.
Habitat:—Usually epiphytic in cloud forest and montane forest at mid to high altitude, elev. 1450–3100 m.
Notes:—The flat transversely oriented leaves remind somewhat of the condition in Phlegmariurus taxifolius;
however in the latter species the stems are usually 2 mm or more thick, and the leaves more closely set and borne in
alternating whorls of 3 or 4. In P. linifolius var. planifolius the leaves are most often inserted singly or in pairs (not
opposite, but at an angle of ca. 120°) alternating with singly inserted leaves.
Specimens examined:—César: Sierra Nevada de Santa Marta, SE slopes, along Río Donachui between Sogrome
and Sacarecungüe, 1742 m, Plowman & Davis 3669 (GH). Magdalena: Sierra de Perijá, 6 km ENE of Manaure, 42 km
E of Valledupar, 7 km from Venezuelan border, 2100 m, Grant 10749 (US). Santander del Norte: Ocaña, 2440–3050
m, Schlim 470 (BR), 2440 m, Schlim 1031 (BR).
FIGURE 18. A. Phlegmariurus linifolius var. jenmanii, (Ecuador, Brandbyge & Azanza 30596, AAU), growth habit. A1. Close-up of
basal division. A2. Close-up of terminal sporangiate division. Del. K. Tind.—B. Phlegmariurus sarmentosus (Ecuador, Holm-Nielsen et
al. 3617, AAU), growth habit. B1. Close-up of basal division. B2. Close-up of sporangiate division.—C. Phlegmariurus sarmentosus
(Camp E-4445, AAU), growth habit. C1. Close-up of sporangiate division. C2. Close-up of sporophyll. Del. B. Johnsen
Phlegmariurus sarmentosus (Spring) B.Øllgaard (2012b: 18).—Fig. 18B.
Lycopodium sarmentosum Spring (1849: 13).—Urostachys sarmentosus (Spring) Herter (1923: 165).—Huperzia sarmentosa (Spring)
Trevisan (1874: 248). Type:—ECUADOR. Pichincha (?): Quitonian Andes, herb. Hooker, Jameson 41 (lectotype K, designated by
Øllgaard (1988: 100).
Illustrations: Øllgaard (1988: Fig. 15D).
Description:—Plants slender, pendulous, with flaccidly hanging divisions, to 100–200 cm long. Shoots homophyllous
or almost so, equally thick throughout, 15–25 mm in diameter including leaves in basal divisions, or sometimes tapering

to ca. 8 mm in diameter. Stems excluding leaves 1–1.5 mm thick at the base, often tapering to 0.4–0.8 mm, slightly to
prominently ridged by decurrent leaf bases, discontinuously or ultimately continuously sporangiate from 10–30 cm
above the base and upward, to 7(–11) times dichotomous. Leaves borne in alternating, irregular, often oblique, whorls
of 4–7, these (1–)1.5–2(–3) mm apart, forming 8–14 obscure longitudinal ranks, spreading to ascending, or rarely
loosely appressed in terminal divisions, usually twisted at the lamina base, straight to slightly upward curved, linear
to linear-subulate, with distinctly widened, auriculate, usually strongly revolute, often overlapping lamina bases, 7–15
× 0.4–0.7 mm just above the auriculate base, usually ca. 0.3 mm wide at the middle, usually somewhat revolute, with
obscure to prominent vein, with smooth margins, or the auricles irregularly toothed. Leaves of terminal divisions often
densely crowded, conform, or gradually reduced to 5–7 mm long. Decurrent leaf bases narrower than the lamina base.
Sporangia 1–1.3 mm wide.
Distribution:—Colombia, Ecuador and Northern Peru.
Habitat:—Epiphytic or rarely hanging from banks, in upper montane forest, elev. 2500–3100 m.
Notes:—Phlegmariurus sarmentosus is quite variable with respect to leaf length. It resembles P. mollicomus,
P. polycarpos, and pendulous forms of P. wilsonii somewhat, but differs from these by its auriculate lamina bases.
Phlegmariurus watsonianus Maxon (1912: 29) with type from Panama, is closely related.
Specimens examined:—Antioquia: Urrao, páramo de Frontino, El Río, 3050 m, Londoño et al 678 (AAU, HUA,
MEDEL). Medellín, Henri-Stanislas, Bro. 1706 (US). Chocó: Mun. San José de Palmar, Cerro de Torrá, E slope, 2500
m, Silverstone-Sopkin et al. 4342, 4923 (AAU). Province unknown: Mutis 3354 (US).
Group of Phlegmariurus brongniartii
Phlegmariurus brongniartii (Spring) B.Øllgaard (2012b: 12).—Fig. 9B.
Lycopodium brongniartii Spring (1841: 515).—Huperzia brongniartii (Spring) Trevisan (1874: 248).—Lycopodium taxifolium Swartz var.
brongniartii (Spring) Baker (1887: 16).—Urostachys brongniartii (Spring) Nessel (1927: 388). Type:—BOLIVIA. La Paz: Yungas,
D’Orbigny 227 (holotype P; isotypes BONN-Nessel 169, BR).
Illustrations:—Øllgaard (1988: Fig. 3A).
Description:—Plants erect from an ascending base, to 50 cm tall, sparsely branched, to 3 times dichotomous. Shoots
homophyllous, almost equally thick throughout, 15–25 mm in diameter including leaves. Stems excluding leaves
2.5–4 mm thick near the base, sometimes tapering to 1.5–2 mm upward, somewhat ridged by decurrent leaf bases,
sporangiate from 15–30 cm above the base and upward. Leaves uniform throughout, borne in alternating, often oblique
whorls of 3–4(–5), these 2–6 mm apart, forming 6–10 often indistinct longitudinal ranks, wide-spreading to somewhat
reflexed, usually straight, not twisted at the base, widely lanceolate to elliptic-lanceolate, rounded at base, with acute
apex, papery to subcoriaceous and opaque, 8–12(–15) × (2–)2.5–4 mm, almost flat, with prominent vein above, or
folded slightly down along the vein, with revolute, minutely rugose and, apically, denticulate or ciliolate margins.
Sporangia (1.5–)2(–2.5) mm wide.
Distribution:—Colombia to Bolivia.
Habitat:—Terrestrial in upper montane forest and on road banks and land slides, 2200–3000 m.
Notes:—A heterogeneous species with slightly distinct populations in Colombia-Ecuador and Peru-Bolivia, the
northern population often with wider leaves, and finely denticulate margins of the leaf apices. The minutely rugulate leaf
margins, characteristic of this species, occur also in Phlegmariurus rosenstockiana and may indicate a relationship.
Specimens examined:—Antioquia: Road San José de la Montaña–Toledo, near Quebrada Santa Inés bridge,
2720 m, Santa 472 (US). Mun. Yarumal, Alto de Ventanas, 3000 m, Santa 608 (AAU). Páramo de Sonson, road
Sonson–Nariño, 2750–2925 m, Core 738 (US). Caquetá: Pass between Garzón and Florencia, E slope of Cord.
Oriental, 2000 m, Mason 13950 (UC, US). Cauca: Road Timbío–Veinte de Julio, km 54, 2100–2300 m, Maas &
Plowman 2121 (AAU, GH, U). Mun. El Tambo, road to the sea through 20 de Julio, W slope, 2320 m, Santa et al. 988
(AAU). Chocó: Mun. San José de Palmar, Cerro de Torrá, summit, 2700–2800 m, Silverstone-Sopkin 4651 (AAU).
Meta: Cord. Oriental, Rio del Nevado, E end of Cordillera de las Cruces, S of Río Arroz, across from El Capitolis,
prominent peak in Cord. Los Laureles, Sumapaz region, 2675 m, Fosberg 20900 (US). Nariño: Mun. Guachucal,
Páramo de Infernillo, 3250 m, Stancik 2633 (PSO). Putumayo: Road Mocoa–Sibundoy, km 112–113, 2440 m, Luteyn
et al. 5072 (COL, GH, NY, UC, US). Road Pasto–Mocoa, km 140, 2200 m, Maguire 61821 (NY).
FIGURE 19. Phlegmariurus linifolius var. planifolius (Funck & Schlim 984, BR). A. Leaves of basal division. B. Terminal sporangiate
division.
Phlegmariurus rosenstockianus (Herter) B.Øllgaard (2012b: 18).—Fig. 15C-D.
Lycopodium rosenstockianum Herter (1909b: 90).—Urostachys rosenstockianus (Herter) Nessel (1939: 174).—Huperzia rosenstockiana
(Herter) Holub (1985: 76). Type:—ECUADOR: Ost-Abhang des Tungurahua, 3000 m, Rimbach 130 (holotype Herb. Rosenstock,
not located; isotypes S, UC fragment, US).
Urostachys cassandrae Herter (1954: 116).—Lycopodium cassandrae (Herter) Morton (1964: 72). Type:—ECUADOR..Pichincha/Napo:
Along trail to Camp Arellan, E of Cayambe, 10300 ft., Drew E-344 (holotype US; isotype MSC).
Urostachys socratis Herter (1954: 117).—Lycopodium socratis (Herter) Morton (1964: 72).—Huperzia socratis (Herter) Rolleri &
Deferrari (1988: 157). Type:—COLOMBIA: Santander, W slope of Páramo Rico, E Cordillera, 3300–3600 m, Killip & Smith 17749
(holotype US; isotype GH)
Illustrations: Øllgaard (1988: 14C, 14D).
Description:—Plants lax, pendulous, rarely recurved from an ascending base, at least to 150 cm long. Shoots
homophyllous, or with gradually slightly smaller leaves upward, 20–25(–30) mm in diameter including the leaves in
basal divisions, sometimes tapering to 10–14 mm in diameter in terminal densely sporangiate divisions. Stems excluding
leaves (1.5–)2–3(–5) mm thick at the base, tapering to 1–2 mm upward, pale greenish to brownish, sporangiate from
30–70 cm above the base and upward, usually 4–7 times dichotomous. Leaves of basal divisions borne in alternating
whorls or irregular low spirals of 3–5, these 3–6 mm apart, forming 6–10 indistinct longitudinal ranks, irregularly
spreading to ascending, usually somewhat twisted at the base, with oblique to vertical lamina, lanceolate to narrowly

oblong, widest at or below the middle, with broadly cuneate to somewhat rounded base, with short to long acute apex,
widely attached to the stem, 10–15 × 2–4 mm, subcoriaceous and opaque to herbaceous and translucent, almost flat,
with prominent vein above, or sometimes folded down along the vein, with slightly revolute, minutely rugose margins
with individually protruding epidermis cells, especially at the apex. Leaves of middle and terminal divisions conform,
or gradually smaller and more ascending to loosely appressed, more densely crowded, the whorls usually 1.5–2.5 mm
apart. Leaves of densely sporangiate divisions usually widest just above the base, lanceolate to narrowly triangular,
with a rounded base, 6–9 × 1.5–2.5 mm, partly covering the sporangia. Sporangia 1.5–2 mm wide.
Distribution:—Colombia, Ecuador, Peru.
Habitat:—Epiphytic, or sometimes terrestrial, in cloud forest and elfin forest, elev. 2400–3500 m. In sheltered
sites in very humid forest at the timberline.
Notes:—Phlegmariurus rosenstockianus seems most closely related to the erect-growing P. brongniartii with which
it shares the details of the rugulate leaf margins. Some Ecuadorian collections seem intermediate between the two
species, but are placed here because of the recurved tips of the erect shoots, and the narrowly oblong rather than
lanceolate leaves.
Specimens examined:—Antioquia: Mun. Urrao, Páramo de Frontino, Llano Grande, 3320–3450 m, MacDougal
et al. 4464 (MO, UC). Cauca: Cord. Central, Quebrada Manchai, NE of Silvia, 3200 m, Haught 5115 (COL, US).
Paletará, Cord. Central, forest E of Llano, 3100–3300 m, Pennell 7013 (GH, K, NY, S, US). Volcán Puracé, 3000
m, Yepes Agredo et al. 18121 (HUA). Macizo Colombiano, Valle de Las Papas, near Valencia, 2910 m, Idrobo et al.
3774 (COL, G). Páramo de Puracé, W slopes of valley of Río Cocuy, 3400 m, Killip & Lehmann V. 38606 (COL, US).
Puracé, Cord. Central, 3450 m, Sneidern 1696 (S). Cundinamarca: 8 km W of Caqueza, 3100 m, Barkley & Mullen
38CO48 (GH). Nariño: Drainage of upper Tescual, above Córdoba, 2840–2890 m, Ewan 16294 (BM, COL, GH,
NO, S, UC, US). Norte de Santander: Ocaña, 2440–3050 m, Schlim 506 (BR, G, K, P), 642 (BR, G, P). Province
unknown: Mutis 3362 (NY, US). Moritz 231 b (G).
Group of Phlegmariurus reflexus
This is a group of terrestrial species, mainly pioneers on disturbed ground, like road banks and land slides in montane
forest.
Phlegmariurus acifolius (Rolleri) B.Øllgaard (2012b: 11).—Fig. 21.
Lycopodium acifolium Rolleri (1985: 2).—Huperzia acifolia (Rolleri) Rolleri & Deferrari (1988: 155). Type:—PERU. Ayacucho: Cearrapa,
between Huanta and Río Apurimac, 1500 m, Killip & Smith 22368 (holotype US).
Description:—Erect, or erect from a decumbent base, 15–30 cm tall, or to 60 cm long. Shoots homophyllous, almost
equally thick throughout, 10–20 mm in diameter including leaves, shoot apices erect. Stems excluding leaves 2–5(–6)
mm thick at the base, sometimes tapering to 1–3 mm thick. Leaves uniform throughout, borne in alternating, usually
densely crowded whorls of 8–11, usually sharply bent upward from a perpendicular junction to the stem and then
gently to strongly claw-like recurved, linear-subulate, widest just above the pale and soft-herbaceous base, 6–10 ×
0.7–1(–1.3) mm, narrowed shortly above the base, subcoriaceous at apex, adaxially convex distally, abaxially flat to
convex with obscure to somewhat prominent vein (sometimes sunken when dried), with rather densely denticulate-
ciliolate margins at base, usually sparsely denticulate or smooth at apex. Sporangia 1–1.5 mm wide.
Distribution:—Colombia to Peru.
Habitat:—Moist banks in montane forest at mid-altitudes, elev. 1400–2300 m.
Notes:—Related to Phlegmariurus reflexus and P. unguiculatus.
Specimens examined:—Cauca: El Tambo, road to the sea through Veinte de Julio, 1850 m, Santa et al. 995
(AAU, COL, HUA). Huila: Cord. Oriental, E of Neiva, 1800–2300 m, Rusby & Pennell 882 (NY). Road Pitalito–
Mocoa, km 28–32, near divide, 2100–2240 m, Luteyn et al. 7549 (AAU, NY). Putumayo: Río Susunga, upper Río
Mocoa, 18–20 km W of Mocoa, 1875 m, Fosberg 20394 (US, p.p.).
FIGURE 20. A1. Phlegmariurus reflexus (Ecuador (Holm-Nielsen et al. 3873, AAU) growth habit. A2. Close-up of sporangiate
division.—A3. Phlegmariurus reflexus (Holm-Nielsen et al 207, AAU) Close-up of sporangiate division. —B1. Phlegmariurus affinis
(Ecuador, Asplund 16257, S), growth habit. B2. Close-up of sporangiate division. B3. (Asplund 16271, S), Close-up of sporangiate
division.—C1. Phlegmariurus eversus (Ecuador, Holm-Nielsen 5644, AAU), growth habit. C2. Close-up of sporangiate division.—C3.
Phlegmariurus eversus (Ecuador, Holmgren 674, S), close-up of sporangiate division. Del. B. Johnsen.

Phlegmariurus affinis (Trevisan) B.Øllgaard (2012b: 11).—Fig. 20B.
Lycopodium affine Greville & Hooker (1831: 364), non Bory (1804: 204).—Huperzia affinis Trevisan (1874: 248).—Lycopodium
blepharodes Maxon (1914: 423).—Urostachys affinis (Greville & Hooker) Nessel 1935: 69).—Urostachys blepharodes (Maxon)
Herter (1949a: 53).—Huperzia blepharodes (Maxon) Holub (1985: 71).—Lectotype:—ECUADOR. Pichincha, 1824, Herb. Greville,
Jameson s. n. (lectotype K, designated by Nessel (1939: 97), possible isotypes E, NY, US).
Urostachys involutus Nessel (1940a: 162).—Lectotype:—ECUADOR: ad radices montis Pichincha, Jameson s. n. (lectotype BONN,
Herb. Nessel 180 p. p., designated by Øllgaard (1988: 30).
Description:—Plants erect from a decumbent base, to 25 cm tall (–60 cm long), to 5 times dichotomous. Shoots
homophyllous, equally thick throughout, 8–15 mm in diameter including leaves. Stems excluding leaves 2–3 mm
thick at the base, upward tapering, sporangiate almost from the base. Leaves borne in alternating whorls of 5, these
1.5–3 mm apart, subdistant, forming 10 longitudinal ranks, usually not covering the stem, ascending to reflexed,
narrowly triangular-lanceolate to almost subulate, with slightly widened base, 6–8 × 1.2–2 mm, abaxially convex with
prominent veinal ridge, at least at base, somewhat clasping the sporangia, with long-ciliate margins. Sporangia 1.5–2
mm wide.
Distribution:—Colombia to Peru
Habitat:—Terrestrial on sheltered road banks in upper montane forest, in Colombia known from the western
Andean slopes, elev. 1800–2800 m. Ramírez Padilla & Macias Pinto (2007) reported this species from Dept. Cauca.
Notes:—Phlegmariurus affinis is variable with respect to crowding and direction of the leaves. It often grows
together with P. reflexus and P. eversus. It is easily distinguished from these by its wider leaf bases, and ciliate leaf
margins.
While Lycopodium affine Greville & Hooker is illegitimate, Trevisan’s combination of the epithet with Huperzia
is valid according to the Code (Art. 72.1) when treated as a new name.
Specimens examined:—Nariño: Mun. Tuquerres, road Balalaika–Yascual, 1800–2500 m, Mora 373 (COL p.p.);
Pass W of Tuquerres, 2800 m, Alston 8414 (AAU).
Phlegmariurus eversus (Poiret) B.Øllgaard (2012b: 15).—Fig. 20C.
Lycopodium eversum Poiret (1814: 556).—Lycopodium reflexum Willdenow (1810: 52), non Lamarck (1792: 653).—Huperzia eversa
(Poiret) B.Øllgaard (1988: 28). Type:—ECUADOR: (Humboldt) Nee D. D. s. n.; Tungurahua (holotype B-Willdenow 19419).
Lycopodium reflexum Lamarck var. polycarpum Sodiro (1883: 90).—Lycopodium polycarpum (Sodiro) Underwood & F. Lloyd 1906:
105), non L. polycarpos Kunze (1835: 5). Type:—ECUADOR: Ecuador, valle de Nanegal, Sodiro s. n.; not located.
Lycopodium ecuadoricum Herter (1909a: 48).—Urostachys ecuadoricus (Herter) Herter (1949: 59).—Huperzia ecuadorica (Herter) Holub
(1985: 72). Type:—ECUADOR: Andium nemoribus humidis, Jameson 74 (lectotype P, designated by B.Øllgaard (1988: 28).
Urostachys dingesianus Nessel (1935: 69). Type:—ECUADOR: Quito, Jameson s. n. (holotype BONN-Nessel 183a).
Urostachys ringshausenii Nessel (1935: 68–69). Type:—COLOMBIA: “Dept. of El Cauca, Shrub zone (Paramilla), 1700–3000 m, Jun.
30, 1922” Collector unknown (holotype BONN-Nessel 154a).
Description:—Plants erect or erect from a decumbent base, soft, often forming large rich-branched clumps, to 30(–50)
cm tall, to 8 times dichotomous. Shoots homophyllous, equally thick throughout, (2.5–)3–6(–10) mm in diameter
including leaves. Stems excluding leaves 1.5–2.5 mm thick at the base, sometimes tapering to 1–2 mm, prominently
ridged by decurrent leaf bases (dried), sporangiate from 5–15 cm above the base and upward. Leaves uniform throughout,
borne in often irregular and oblique, alternating, subdistant to densely crowded whorls of (4–)5(–6), these 1–4 mm
apart, forming (8–)10(–12) longitudinal ranks, wide-spreading to sharply reflexed, usually strongly recurved, linear-
lanceolate, widest in the basal half, (2.5–)3–5 × 0.5–1 mm, softly herbaceous to subcoriaceous, adaxially convex with
obscure vein, abaxially irregularly concave (dried), with obscure to somewhat prominent vein, with slightly revolute,
denticulate-ciliolate margins. Leaf base with prominently decurrent vein and margins. Sporangia 1–1.5 mm wide.
Distribution:—Costa Rica, Panamá, Andes from Venezuela to Bolivia.
Habitat:—Terrestrial, as a pioneer on landslides, road banks, and other open, moist habitats in upper montane
forest, elev. 1500–3500 m.
FIGURE 21. A. Phlegmariurus unguiculatus (Colombia: Schwabe 169/071, COL), growth habit. A1. Close-up of terminal fertile
division.—B. Phlegmariurus acifolius (Colombia: Luteyn 7549, AAU), growth habit. B1. Close-up of terminal fertile division.
Notes:—Phlegmariurus eversus resembles P. reflexus, but differs by usually forming very densely branched
individuals, and by having shorter and relatively wider leaves, which are usually strongly recurved, usually fewer
together in each whorl, and leaving the stem more exposed.
Urostachys rolandii-principis Nessel (1939: 117): Identity uncertain as no type was designated, but some of
the collections cited belong to Phlegmariurus eversus, and one of them is the lectotype collection of Lycopodium
ecuadoricum Herter, cited above as a synonym of P. eversus.
Specimens examined:—Antioquia: Urrao, road to páramo de Frontino, ‘La Ilusión’, 3000 m, Santa 1173 (AAU,
COL, HUA); Ibid., 3150 m, Santa 1174 (AAU). Urrao, trail to Páramo de Frontino, 3000–3300 m, McPherson 13205
(AAU). Mun. Urrao, Jaiperá, Vereda El Chuscal, Páramo de Frontino, Llanogrande, 3000–3390 m, Callejas et al.
7453, 7538 (AAU). S of Medellín, SE of Caldas, Alto de Morrogil, 2450–2560 m, Juncosa & Misas 1103 (AAU, MO).
Andes Centr. Antioquia, 2000–3000 m, Mayor 44 (S). Bolívar: Below Páramo de Chaquiro, Cord. Occ., 2800–3100
m, Pennell 4292 (F, GH, K, US). Boyacá: N of Belán, 10100 ft, Fassett 25903 (US). Caldas: Nevado del Ruiz,
Road Manizales–Termales, 3250 m, Cleef & Florschütz 6021 (AAU). Nevado del Ruiz, road Manizales–Termales,
3000–3500 m, Forero et al. 550 (COL, F, US). Cord. Central, W slope, road to Termales, near La Enea, 2700–3000 m,
Humbert et al. 27007 (HBG, P, US). Nevado del Ruiz, Road Manizales–Termales, 3080 m, Santa & Hoyos 676 (AAU,
QCA). Páramo del Ruiz, 25 km SE of Manizales, 3400 m, Tryon & Tryon 6149 (COL, GH, K, NY, US). Manizales–
Nevado del Ruiz, km 13. 2500 m, Bischler 1470 (G). Caldas/ Tolima: Quindío pass, 3300 m, Grant & Drew 10616
(US). Cauca: Cord. Occ. Los Farallones, extreme N, E slopes, Alto de Buey, 3300–3450 m, Cuatrecasas 18065 (F, S,
US). Cordillera occidental, Los Farallones, Alto del Buey–Quebrada de Los Ramos, 3450–3350 m, Cuatrecasas 18029
(F, US). Alto de Achupallas, between Almaquer and La Cruz, 3000 m, Lehmann 6221 (K, US). Páramo de Puracé, 8
km E of Puracé, 25 km E of Popoyan, 3000–3300 m, Tryon & Tryon 5975 (COL, GH, K, NY, S, US), 5976 (COL,
GH, NY, US). Near Pico de Aguila, W of Morales, 2750 m, Core 1104 (GH, MO, US). Macizo Colombiano, Valle
de las Papas, alrededores de Valecia, 2910 m, Idrobo, Pinto & Bischler 3786 (G). Alto Todos Santos–San Sebastian,
2300 m, Core 1064 (US). Andes of Popoyan, Lehmann 6970 (K, US). Totoró–Inzá, E slope of Cord. Central, between
Páramo de Las Delicias and Inzá, 2980 m, Santa et al. 985 (AAU). Parque Nacional Puracé, Paletará division, km

37–43, 3050–3150 m, Luteyn 6938 (AAU, NY). Volcán Puracé, 3280 m, Cleef 2675 (AAU). Puracé, 3180 m, Santa &
Hoyos 716 (AAU, QCA). Cundinamarca: Cord. Oriental, summit of Cordillera de Heliconia, 13 km SE of Gachalá,
15 km NW of Medina, 2930 m, Grant 10250 (COL, US). Mun. Bogotá, road to Choachí, 3100 m, Santa & Escobar
1128 (AAU). Huila: La Plata, between San Augustin and Popoyan, 1525 m, Hey C2 (U). Nariño: Above El Encano,
3200 m, Hernández 16 (US). Mun. Pasto, Road to páramo de Quillinsayaco, 3060 m, Santa & Buitrago 922 (AAU,
COL, HUA). Above Río de Potreros, headwaters of Río Tellez, 3200–3400 m, Ewan 16529 (GH, NO, UC, US). Cerro
Macha, W side of Volcán Doña Juana, Cord. Oriental, 3200 m, Ewan 16606 (GH, NO, US p. p.). Laguna Cocha,
André 2995 (K). Mun. Ricaurte, La Planada, ca. 5 km S of Chucunés, 1850 m, Luteyn 6832 p.p. (AAU, COL, NY).
Tabano, near Pasto, André K 1466 (K). Mun. Pasto, La Laguna near Pasto, Barclay et al. 243 (AAU). Putumayo:
Comm. del Putumayo, 2960 m, Fosberg 20437 (US). Beyond Sibundoy at San Francisco, 2260 m, Foster 1975 (GH).
Road San Francisco–Mocoa, 2300 m, Bristol 411 (GH, US). Risaralda: Cerro Tatamá, 3400–3700 m, Pennell 10543
(US). Camino Termales de S. Rosa de Cabal—La Sierra. Finca Berlín, 3000 m, Boots 17 (AAU). Santander Norte:
Pamplona, Garganta de 945 (F). Tolima: Boqueron de Quindiu, 3400 m, Alston 7722 (AAU, BM). Boqueron de
Quindiu, André 2245 (K, NY). Ibagué, trail to nevado del Tolima, 2760 m, Santa et al 963 (AAU, COL, HUA).
Quindío Pass, 35 km W of Ibague on road to Armenia, 3100–3200 m, Tryon & Tryon 6022 (COL, GH). Valle del
Cauca: Alto del Pan de Azucar, road Tenerife–Pan de Azucar, 3000 m, van Rooden et al. 481 (AAU, Z).
Phlegmariurus luteynii B.Øllgaard (2015: 194–195).—Fig. 22.
Type:—COLOMBIA. Valle/Chocó: Valle/Chocó border, Mun. El Cairo, Correg. Boquerón, Vereda Las Amarillas, Serrania Las Paraguas,
ca. 04°45’N 76°20’W, along road to and beyond Cerro del Inglés, 17–23 km W of El Cairo, 1750–2050 m,Luteyn, Silverstone-
Sopkin, Heredia & Paz 12304 (holotype NY; isotype CUVC).
Illustrations: Øllgaard (2014: Fig. 10).
Description:—Plants erect, to 25 cm tall, sparsely branched, to 4 times dichotomous. Shoots homophyllous or the leaves
slightly smaller in terminal divisions, almost equally thick throughout, 10–15 mm in diameter including leaves. Stems
excluding leaves 1–2(–2.5) mm thick near the base (dried), sometimes tapering to 1 mm upward, somewhat ridged by
decurrent leaf bases, usually distinctly red, sporangiate from 1.5–7 cm above the base and upward. Leaves uniform
throughout, borne in alternating, often oblique whorls of 4, these 1.5–2 mm apart, forming 8 indistinct longitudinal
ranks, wide-spreading, usually straight, not twisted at the base, lanceolate to elliptic, in the lower divisions often
somewhat attenuate at the base, with acute apex, papery and slightly translucent, 5–7.5 mm long in basal divisions,
upward gradually 3.5–5 mm long, (1.5–)2–2.5 mm wide, almost flat or with slightly revolute margins, with distinct but
not prominent vein, the margins distinctly and densely ciliate. Sporangia 1.5–2 mm wide. Spores normal , ca. 32 μm.
Distribution:—Endemic to Colombia, Deptos El Chocó and El Valle del Cauca.
Habitat:—Apparently a pioneer species on road banks.
Notes:—Recognized by the red stems and densely ciliate leaf margins. The position of this species in this group
is uncertain.
Specimens examined:—Valle del Cauca: Ansermanuevo-San José del Palmar road, km 60–62, steep roadside
slopes and thickets, 1870–1950 m, Luteyn, Lebrón-Luteyn & Morales 7290 (COL). Anchicayá (Cali–Buenaventura)
500 m, Køie 4771 (C).
Phlegmariurus reflexus (Lamarck) B.Øllgaard (2012a: 481).—Fig. 20A.
Lycopodium reflexum Lamarck (1792: 653).—Plananthus reflexus (Lamarck) Palisot de Beauvois (1805: 100).—Urostachys reflexus
(Lamarck) Herter (1922: 249).—Huperzia reflexa (Lamarck) Trevisan (1874: 248). Type:—MARTINIQUE: herb. Lamarck, Comm.
Joseph Martin s.n. (holotype P).
Lycopodium bifidum Willdenow (1810: 53).—Lycopodium reflexum Lamarck var. majus Spring (1842: 26).—Huperzia reflexa (Lamarck)
Trevisan var. bifida (Willdenow) Trevisan (1874: 248).—Urostachys bifidus (Willdenow) Nessel (1939: 110).—Huperzia bifida
(Willdenow) Holub (1985: 71). Type:—VENEZUELA: Cuchilla de Guajana Guajana, Humboldt & Bonpland 474 (holotype B-
Willdenow 19421; isotype P-Humboldt).
Lycopodium reversum Presl (1825: 82).—Urostachys reversus (Presl) Herter (1949: 78). Type:—ECUADOR. Guayaquil: Guayaquil,
Haenke s. n. (holotype PRC).
Lycopodium reflexum Lamarck var. densifolium Baker (1887: 11).—Lycopodium densifolium (Baker) Underwood & Lloyd (1906: 106).—
Urostachys reflexus (Lamarck) Herter var. densifolius (Baker) Nessel (1927: 394).—Urostachys densifolius (Underwood & Lloyd)
Herter (1949: 58). Syntypes:—COLOMBIA: Popayan, Hartweg 1480 (G, K); Moritz 2266 (=226b: HBG, K); BRAZIL: Glaziou
15797 (K).
Lycopodium pearcei Baker (1887: 14).—Urostachys pearcei (Baker) Nessel (1939: 73).—Huperzia pearcei (Baker) Holub (1985: 75).
Type:—BOLIVIA. Santa Cruz: Feb., 1864 (holotype K; isotype BONN-herb. Nessel 116).
Urostachys jergii Nessel (1935: 70). Type:—PUERTO RICO: Sintenis 6345 (holotype BONN-Nessel 229; isotype NY).
Lycopodium mexiae Copeland (1941: 294).—Huperzia mexiae (Copeland) Rolleri and Deferrari (1988: 156). Type:—PERU: Huánuco:
Churubamba, trail Cotirarda–Mercedes, 1875 m, Mexia 8193 a (holotype UC; isotypes F, GH, K, MICH).
Urostachys leptodon Herter (1954: 120–121).—Lycopodium leptodon (Herter) Morton (1964: 72).—Huperzia leptodon (Herter) Rolleri &
Deferrari (1988: 156). Type:—COLOMBIA: Guasca, Bro. Ariste-Joseph A132 (holotype US 888340).
Urostachys stellae-polaris Herter (1954: 121).—Lycopodium stellae-polaris (Herter) Morton (1964: 72). Type:—COLOMBIA.
Cundinamarca: Guayabetal to Monte Redondo, SE of Quetame, 1300–1500 m, Pennell 1801 (holotype US; isotypes GH, NY).
Lycopodium spongiosum Rolleri (1985: 6).—Huperzia spongiosa (Rolleri) Rolleri & Deferrari (1988: 157). Type:—COLOMBIA:
Cundinamarca: Chocontá–Guateque highway, 2900 m, Haught 5906 (holotype US, isotypes COL, F, NY, S)
Illustrations: Øllgaard (1988: Fig. 4A); Mickel & Beitel (1988: Fig. 1E, F); Lellinger (1989: Fig. 36).
FIGURE 22. Phlegmariurus luteynii (Luteyn et al. 12304 (NY). A. Growth habit of large and small plants. A1. Close-up of terminal fertile
division. A2. Close-up of leaf showing margin character.

Description:—Plants erect or ascending from a decumbent base, soft, usually forming small loose clumps, 10–30(–
40) cm tall. Shoots homophyllous, almost equally thick throughout, 7–15 mm in diameter including leaves. Stems
excluding leaves 1.5–3(–4) mm thick at base, sometimes tapering to 1–1.5 mm thick, ridged by decurrent leaves or
almost smooth, sporangiate from 2–12 cm above the base and upward, usually 2–5 times dichotomous. Leaves borne
in alternating, often irregular and oblique, whorls of (6–)7–8(–9), these 0.7–2.5 mm apart, forming 12–18 longitudinal
ranks, ascending to spreading or sharply reflexed, straight to strongly recurved, linear-subulate, widest just above the
base, 4–8 × 0.5–1(–1.2) mm, softly herbaceous to subcoriaceous, adaxially convex, or concave near the base, abaxially
flat, or slightly concave to convex, with obscure to somewhat prominent vein, with flat to revolute, very sparsely to
densely denticulate to short-ciliolate margins. Leaf bases often somewhat decurrent. Sporangia 1–1.5 mm wide.
Distribution:—Throughout humid mountainous regions of Tropical America.
Habitat:—Terrestrial, erect or ascending herbs. Road banks and other open, humid situations, generally in
montane forest, elev. 370–3400 m.
Notes:—Phlegmariurus reflexus is widespread in the Neotropics. It is a frequent pioneer on permanently moist,
disturbed ground, but may also occur in peat bogs, or even, by chance, as a low epiphyte. It is closely related to P.
eversus, P. affinis, P. acifolius, and P. unguiculatus. Phlegmariurus firmus seems more distantly related.
Phlegmariurus reflexus is highly variable with respect to leaf and stem size, direction and crowding of leaves,
and leaf margin characters. Part of the variation may reflect growth conditions, but often several different forms may
be found growing intermixed in the same habitat, indicating that genetic differences exist. The variation patterns are
complex and in need of a detailed study. The different morphological forms of P. reflexus, as well as other closely
related species, often occur together on large road banks or land slides, and here may be compared under uniform
growth conditions. In such populations the individuals may exhibit subtle or obvious differences. Neighbouring and
more distant populations may exhibit similar variation and thus taxonomic recognition may be indicated. However,
often the differences are not matched by other individuals, neither in the same nor in other populations, and therefore
the plants may have the character of individual aberrants. Some of these are intermediate between other forms and may
be hybrids, without exhibiting hybrid features such as abortive spores or irregular meioses. The slight differences, for
example of leaf shape and direction, appear especially striking in the genus Phlegmariurus, in which the features are
repeated in hundreds of leaves on a stem, creating regular and recognizable patterns. At the same time these features,
by which the individuals differ, appear to be without adaptive significance.
A variety of small dimensions stands somewhat apart, and was earlier recognized taxonomically, but this idea has
been abandoned because it is an exaggerated simplification of the variation.
Specimens examined:—Antioquia: Near Santa Rosa, Barkley 43007 (GH). Near Medellín, Charetier 35 (US).
Near Las Palmas on road Medellín–Retiro, 2500 m, Hodge 6653 (GH). Medellín Valley at Primavera, near Caldas,
1700–1800 m, Hodge 6664 (GH). Road Medellín–Pto. Triunfo, at Cocorna, 2090 m, Juncosa 968 (MO). Road
Medellín–Río Negro, 1900 m, Killip et al. 39823 (COL, US). And. Centr. Antioquia, Mayor 77 (S). Yarumal, 2360 m,
Santa 600 (AAU, COL, HUA). Mun. Urrao, road to Páramo de Frontino, 3250 m, Santa 1175 (AAU). Angelópolis,
1950 m, Barkley & Guiterrez 1712 (COL). Boyacá: Road Sogamoso–Pajarito, Río Cusiana, 2000 m, Bischler 1727
(G). Moniquirá, 1580 m, Cuatrecasas 8137A (COL, F). Valle Río Pomera, 13 km NW of Arabuco, road cut near top
of canyon, 9200 ft, Fassett 25646 (US). Road to Páramo de La Rusia, 3000 m, Uribe 4234 (COL, NY). Sierra Nevada
de Cocuy, Bachira–Bocota, 2000 m, Grubb & Guymer P 77 (COL). Caquetá: Florencia, road to Neiva, Sucre, 1120
m, Santa 1245 (AAU, HUA). Road Garzón–La Florencia, E slope of Cord. Or. 1200 m, Mason 13940 (COL, UC,
US). Cord Oriental between Sucre and La Portada, 1200–1350 m, Cuatrecasas 9145 (COL). Cauca: Popayan, 1800
m, Yepes-Agredo 332 (COL, US). Popoyan, Hartweg 1460 (P). Near Piendamor, 1700 m, Alston 7968 (AAU p.p., BM
p.p.). Road Piendamo–Popayan, 1750 m, Haught 5084 (COL, G). Parque Nacional Puracé, 3050–3150 m, Luteyn 6938
(AAU). San Isidro, Puracé, 2200–2500 m, Pennell & Killip 6455 (US). W of Popayan, Cauca Valley Río Sucio to Río
Piedras, 1500–1700 m, Pennell & Killip 8222 (GH, US). Cord. Central, Puracé, 3300 m, Sneidern 1694 (S). Mun.
Tambo, road to sea via 20 de Julio, 2280 m, Santa et al. 999 (AAU). Near rim of Canyon of Río Cauca, E of Dinde,
1550 m, Core 1133 (US). Mondomo, 1100 m, Espinal 2716, 2833 (COL). Chocó: El Piñon, 2000 m, Araque-Molina &
Barkley 19Ch104 (COL). Cundinamarca: Road E of Guasca, 3000 m, Haught 5792 (COL, US). Chocontá–Guateque
highway, 2900 m, Haught 5906 (COL, F, NY, S). Above La Vega, 2400 m, Murillo et al. 733 (COL). Near Buenavista,
1500 m, Alston 7525 (AAU, BM). Bogotá, San Francisco, Apollinaire, s.n. (G, P). Near Usme, Aristé-Joseph A 408
(US). W of Bogotá, 2750 m, Barkley & Bouthillette 38922 (GH). Páramo de Guasca, W slope, 2800–3300 m, Garcia-
Barriga & Schultes 13515 (COL). Mun. de Fómeque, km 4 along road to Chingaza, 2150 m, Acosta-Arteaga 460
(COL). Cord. Or. Campo Alegre, 40 km N of Villavicencio, 1310 m, Grant 10468 (US). Chocontá–Guateque highway,
2900 m, Haught 5906 (F, NY, S), 5907 (COL, G, NY). SE of Bogotá, Quebrada San Cristóbal, 2740–2990 m, Little
8160 (US). Bogotá, trail towards Ubaqué, 2900 m, Little 9239 (COL, US). San Cristóbal, Bogotá, Niemeyer 190 (US).
Bogotá, Quebrada del Chicó, 2800 m, Schneider 154 (S). Bogotá, Quetame (Duitama), 1850 m, Triana 696 (COL).
Bogotá, Quetame, 1400 m, Triana, J. s.n. (G, P). Huila: W slope of Cord. Oriental above Guadalupe en Resina, 1900
m, Peres-Arbelaez & Cuatrecasas 8385 (COL). Road Plata Vieja–Popayán, mi 8, 1430 m, Maguire & Maguire 44197
(NY). Road Pitalito–Mocoa, 2100–2240 m, Luteyn et al. 7549 (NY). Altamira–Florencia road, km 15, 1465 m, Luteyn
et al. 4848 (NY). Magdalena: Sierra Perijá–Villanueva, 1800 m, Haught 4528 (COL, US). Sierra Nevada de Santa
Marta, Had Cincinati, 1250–1500 m, Martin 3207 (MO). Santa Marta, Hanbury-Tracy 565 (K). Sierra Nevada de Santa
Marta, road Santa Marta to crest of San Lorenzo, El Campano, 1190 m, Santa & de Escobar 627 (AAU, COL, HUA).
Sierra Nevada de Santa Marta, road toward Cuchilla de San Lorenzo, Estación de Inravisión, 2400 m, Santa & de
Escobar 637, 659 (AAU). Sierra Nevada de Santa Marta, road Santa Marta to crest of San Lorenzo, near INDERENA
station, 2100 m, Santa & Escobar 653 (AAU). San Lorenzo, 2130 m, Mägdefrau 1037 (COL). Meta: Cord. Or. 9 mi
NW of Villavicencio, 985 m, Grant 9219 (US). Nariño: Near San Juan, on Río San Juan, ca. 2 km below Mayasquer,
2130 m, Ewan 16084 (NO, UC, US). Mun. Tuquerres, road Balalaika–Yascual, 1800–2500 m, Mora 373 (COL p.p.).
Mun. Ricaurte, La Planada, ca. 5 km S of Chucunés, 1850 m, Luteyn 6832 p.p. (AAU, COL, NY, US). Norte de
Santander: Mun. Cacota, Pamplona–Chitagá, 2460 m, Santa & Escobar 1104 (AAU, HUA). Ocaña, Schlim 641
(BR). Ocaña, 1850, 1225–1525 m, Schlim 57 (G, K, P). Ocaña, Kalbreyer 511 (K). Putumayo: Cord. Central, 1875
m, Fosberg 20394 (US). Quindío: Salento, 1600–1800 m, Pennell, F.W., Killip & Hazen 8851 (GH, NY, S). Salento,
caserío Boiquía, Quebrada Santa Rita, 1800 m, Santa & Hoyos 706 (AAU, COL, HUA, MEDEL). Santander: Páramo
de Fontibon, Pamplona, 2500 m, Alston 7178 (AAU, BM). Velez, 370 m, Fassett 25073 (US, p.p.). Piedecuesta–Las
Vegas, 2000–2500 m, Killip & Smith 15497 (GH p.p., US p.p.). N slope of Mesa de los Santos, 1000–1500 m, Killip
& Smith 15004 (GH, NY, US). Road Los Curos–La Granja, Mesa de los Santos, 1500 m, Langenheim 3063C (COL,
UC, US). Subpáramo NW of Arcabuco, Cerro Berlín, Río Chuqueque region, 2820 m, Cleef et al. 3522 (AAU, US).
Tolima: Boquerón de Quindíu, 3400 m, Alston 7722 (AAU). Quindío highway, Cajamarca–summit of Divide, 3000
m, Killip & Varela 34638 (COL, GH, US). La Trinidad, Libano, 1100–1300 m, Pennell 3330 (GH, MO, NY). 35 Km
W of Ibagué on road to Armenia, 3100–3200 m, Tryon & Tryon 6022 (COL), 6023 (GH). Valle del Cauca: Near
Piendamor, 1700 m, Alston 7968 (AAU). Cord. Occidental, Los Farallones, extreme North, between Allto de Buey
and Quebrada de los Ramos, 3350–3450 m, Cuatrecasas 18029 (US). Cord. Occidental, Los Farallones, 3300–3450
m, Cuatrecasas 18065 (S). Old road Cali–Buenaventura, before Queremal, 1530 m, Santa & S. Hoyos 729 (AAU).
Old road Cali–Buenaventura, 7 km beyond Queremal, 1200 m, Santa & S. Hoyos 734 (AAU, COL, HUA, MEDEL).
Anchicayá, Cali–Buenaventura, 500 m, Køie 4771 (C). Mun. El Cairo, Las Amarillas, near base of Cerro del Inglés,
Serranía de los Paraguas, 2070–2125 m, Silverstone-Sopkin 3750 (AAU). La Habana, Buga, (Cord. Central), Espinal
2015 (COL).
Phlegmariurus riobambensis (Herter) B.Øllgaard (2012b: 18).—Fig. 23.
Urostachys affinis (Greville & Hooker) Nessel var. riobambensis Nessel (1940a: 162).—Urostachys riobambensis (Nessel) Herter (1949:
79).—Huperzia riobambensis (Herter) B.Øllgaard (1987: 169). Type:—”Sodiro, Ecuador: Riobamba” (holotype BONN-Nessel 179
p. p.), probably a mislabelled specimen, as nothing with the aspect of this species is known from Ecuador.
Urostachys castoris Herter (1954: 111).—Lycopodium castoris (Herter) C. Morton (1964: 72). Type:—COLOMBIA: Cundinamarca,
Guadalupe, Bogotá, 3100–3400 m, Haught 5706 (holotype US 2016777; isotypes COL, F).
Description:—Erect or ascending rather flexible plants, often with tufted growth, to 50 cm long, at least to 5 times
dichotomous. Shoots homophyllous or nearly so, 10–20 mm wide including leaves at the base, sometimes tapering to
10 mm in terminal divisions, sporangiate from 10 cm above the base and upward. Stems 2–3 mm thick excluding leaves
at the base and throughout or sometimes tapering to 1.5 mm in terminal divisions, somewhat ridged by decurrent leaf
bases (dried). Leaves borne in whorls of 5–6, these 1.5–4 mm apart, forming 10–12 rather indistinct longitudinal ranks,
patent-ascending to somewat reflexed, usually straight, linear-lanceolate to triangular-lanceolate with a rounded base,
acute, 5–12 × 1.2–2.2 mm, subcoriaceous, lustrous above, with denticulate margins, flat–slightly involute, abaxially
usually convex, vein obscure. Sporangia 1.5–2 mm wide.
Distribution:—Venezuela, Colombia
Habitat:—Terrestrial on road banks, shrub páramo, wet páramo, open turf, elev. 2500–3800 m.
Notes:—Phlegmariurus riobambensis is usually more robust than P. reflexus, with subcoriaceous leaves, and
often with slightly golly leaf upper sides.

Specimens examined:—Antioquia: Mun. Santa Rosa de Osos, Llanos de Cuivá, 2740 m, Santa 944 (AAU, COL).
Boyacá: Road Chita–Socha, 3230 m, Santa & Escobar 1120 (AAU, COL, HUA). Mun. Aquitania, road Sogamoso–
Pajarito, Páramo de Las Cintas, 3100 m, Santa & Escobar 1123 (AAU). Cauca: El Tambo, 2500 m, von Sneidern, K.
2197 (S). Cundinamarca: Mun. La Calera, ascending to Páramo de Palacio, 3000–3200 m, Santa 1024, 1025, 1026
(AAU). La Calera, páramo de Palacio, 3500–3800 m, Santa 1032 (AAU, COL, HUA, MEDEL). Bogotá–Usme road,
Páramo de Chisacá, km 28–33, 3450–3375 m, Luteyn et al. 7751 (NY). Guadalupe, above Bogotá, 3200 m, Haught
6684 (COL, US). Summit of Monserrate, at E edge of Bogotá, 3200 m, Little 8187 (US). Páramo 10 km E of Guasca,
3350–3500 m, Little 7447 (COL, US). Road Bogota–Choachí, slope toward Choachí, 3140 m, Lozano et al. 747
(COL). Páramo de Choachi, near Bogotá, 3200–3300 m, Pennell 2243 (NY). Chipaque, at the pass, 3250 m, Schneider
719 (S). Páramo de Palacio, 10 km E of La Calera, 20 km NE of Bogotá, 3300–3400 m, Tryon & Tryon 6032 (COL,
GH). Páramo de Guasca, 7 km E of Guasca, 40 km NE of Bogotá, 3000–3100 m, Tryon & Tryon 5930 (COL, GH,
NY, US). Los Gaques, below Páramo de Guasca, 3300 m, Alston 7473 (AAU, BM, MO). Páramo de Guasca, W slope,
3300 m, Cleef 462 (AAU). Norte de Santander: Ocaña, 2440–3050 m, Schlim 467 (BONN herb. Nessel 165, BR, G,
K, P). Mun. Chitagá, Presidente, 3400 m, Santa & Escobar 1107, 1107A (AAU). Santander: W slope of páramo Rico,
3300–3600 m, Killip & Smith 17762 (GH, NY).
FIGURE 23. A. Phlegmariurus riobambensis (Alston 7473 (AAU) growth habit. A1. Close-up of terminal sporangiate division.—A2:
(Santa 944, AAU), Close-up of terminal sporangiate division.
Phlegmariurus unguiculatus (B.Øllg.) B.Øllgaard (2012b: 20).—Fig. 21A.
Huperzia unguiculata B.Øllgaard (1988: 26). Type:—ECUADOR. Morona-Santiago: Road Sevilla de Oro–Amaluza, N of San Pablo,
2300 m, Øllgaard & Balslev 9454 (holotype AAU; isotype QCA).
Description:—Plants erect, or erect from a decumbent base, usually with nodding shoot apices, soft, usually forming
loose clumps, to 4(–5) times dichotomous, 15–30 cm tall, or to 60 cm long. Shoots homophyllous, almost equally
thick throughout, 10–15(–18) mm in diameter including leaves. Stems excluding leaves 2.5–4(–6) mm thick at the
base, sometimes tapering to 1–3 mm thick, ridged by decurrent leaf bases to almost smooth, sporangiate from 8–20
cm above the base and upward. Leaves uniform throughout, borne in alternating, often irregular and oblique, usually
densely crowded whorls of 9–11(–12), these 0.7–1.5(–2) mm apart, forming 18–22(–24) obscure longitudinal ranks,
spreading to reflexed, straight to strongly claw-like recurved, linear-subulate, widest just above the base, 5–8(–10) ×
(0.5–)0.7–1(–1.3) mm, narrowed shortly above the base, subcoriaceous, adaxially convex, abaxially flat to convex
with obscure to deeply sunken vein, usually with revolute, entirely smooth to sparsely denticulate-ciliolate margins,
sometimes with only leaf base margins denticulate. Leaf bases with shortly and indistinctly decurrent margins and
vein, in sporangiate leaves usually sharply bent upward from a perpendicular junction to the stem. Sporangia 1–1.5
mm wide.
Distribution:—Colombia and Ecuador. Ramírez Padilla & Macias Pinto (2007) reported this species from Dept.
Cauca.
Habitat:—Land slides, road banks, and other open, often disturbed habitats in humid montane forest, elev. 1800–
2300 m.
Notes:—Phlegmariurus unguiculatus is related to P. reflexus and P. acifolius, with which it has the general aspect
and habitat in common. The three species are often found growing together. Under such circumstances P. unguiculatus
is recognized by its nodding shoot apices and strongly recurved leaves.
Specimens examined:—Cauca: Km 54 of road Timbío–Veinte de Julio, 2100–2300 m, Maas & Plowman 2122
(COL, GH). Chocó: Road Ansermanuevo–San José del Palmar, km 65–67, 1975–1850 m, Luteyn et al 7304 (AAU,
COL, NY, US). Putumayo: Road San Francisco–Mocoa, 2300 m, Bristol 408 (GH). Pasto–Mocoa, l40 km E of Pasto,
2200 m, Maguire & Maguire 61820 (AAU, NY, UC). El Mirador, 2100 m, Schwabe C169/071 (COL).
Group of Phlegmariurus firmus
Phlegmariurus firmus was included in the P. reflexus clade in Testo et al. (2018) as it was done by Øllgaard (1987,
1992) and shares most morphological characters with that terrestrial group. I treat it here in a separate group due to the
striking features of the ± isodiametric, thick-walled upper leaf epidermis cells in the species listed here.
Phlegmariurus cleefianus (B.Øllg.) B.Øllgaard (2012b: 13).—Fig. 24.
Huperzia cleefiana B.Øllgaard (2004: 34–35). Type:—COLOMBIA: Huila, Parque Nacional Cueva de los Guácharos y Cerro Punta.
flanco huilense del Cerro, 2740 m, Cleef 5033 (holotype AAU, isotype U).
Illustrations: Øllgaard (2004: Fig. 3).
Description:—Plants erect or erect from a decumbent base, robust, to 150 cm tall, sparsely branched. Shoots
homophyllous, equally thick throughout, 10–15 mm in diameter including leaves. Stems excluding leaves 5–6 mm
thick at the base (dried), sometimes tapering to 4 mm, usually not concealed by the leaves, sporangiate from 60–70
cm above the base and upward. Leaves uniform throughout, borne in alternating, often oblique, subdistant crowded
whorls of 7–8, these 1.5–5 mm apart, forming 14–16 longitudinal ranks, recurved to reflexed, linear to subulate, almost
parallel-sided in the basal half or more, short-acute, 8–10 × 1.5–2 mm, coriaceous, flat or adaxially convex, with
obscure to somewhat prominent vein above, abaxially flat to slightly concave, with narrowly to broadly prominent
vein, with flat to slightly revolute, irregularly rugose margins, hypostomatic. Leaf bases with prominently decurrent
margins and vein. Upper leaf epidermis cells ± isodiametric, thick-walled. Sporangia ca. 2 mm wide.
Distribution:—Endemic to Colombia.
Habitat:—This terrestrial species is endemic to perhumid high-andine forest in southern Colombia. All collections
were made at ca. 2750 m elev. With shoots to 1.5 m, this is the tallest terrestrial Phlegmariurus species I have seen, and
it is not very likely to be overlooked in other places. The specimens are fertile only in their uppermost ca. 25 cm.
Notes:—Murillo & Murillo (1999) referred the cited specimens to Huperzia firma (Mett.) Holub (=Phlegmariurus
firmus) to which P. cleefianus seems closely related.
Specimens examined:—Cauca: Cordillera Central, Macizo Colombiano, flanco E. Parque Nacional de Puracé,
subida desde S. José de Isnos. Bosque alto-andino súmamente húmedo con Weinmannia, Clusia, Miconia spp.,
Bromeliáceas epífitas y terrícolas, Geonoma y Sphaeradenia. Planta terricola 1.5 m, asociada con Bromeliáceas

terrestres y Sphagnum y bosque abierto, 2750 m, Cleef 5002 (AAU, U). Cord. Central, Quebrada Manchai, NE of
Silvia, 3200 m, Haught 5117 (COL). Huila: Mun. San José de Isnos, Vereda El Marmol, Parque Nac. de Puracé, 2800
m, Lozano 3420 (COL).
FIGURE 24. Phlegmariurus cleefianus (Cleef 5033, AAU), parts of growth habit. A1. Close-up of terminal sporangiate division.
Phlegmariurus firmus (Mettenius) B.Øllgaard (2012b: 15).—Fig. 10D.
Lycopodium firmum Mettenius (1865: 307).—Urostachys firmus (Mettenius) Nessel (1927: 391).—Huperzia firma (Mettenius) Holub
(1985: 72). Type:—COLOMBIA: Bogotá, La Peña, 2900 p., Goudot, coll. Lindig 1521 (holotype P; isotypes B, K, NY fragm.).
Lycopodium coriaceum Rolleri (1981: 85).—Huperzia coriacea (Rolleri) Rolleri & Deferrari (1988: 156). Type:—COLOMBIA.Caldas:
Cerro Tatama, Cord. Occidental, 3400–3700 m, Pennell 10542 (holotype US).
Illustrations: Øllgaard (1988: Fig. 2D).
Description:—Terrestrial, erect or erect from a decumbent base, robust, usually 10–30 cm tall, sparsely branched, to
4 times dichotomous. Shoots homophyllous, equally thick throughout, 8–15 mm in diameter including leaves. Stems
excluding leaves 3–5 mm thick at the base, sometimes tapering to 2.5–3 mm, often concealed by leaves, sporangiate
from 10–50 cm above the base and upward. Leaves uniform throughout, borne in alternating, often oblique, subdistant
to densely crowded whorls of 7–10, these 1–4 mm apart, forming 14–20 longitudinal ranks, ascending to reflexed,
not or slightly sigmoid, linear to subulate, often almost parallel-sided in the basal half or more, short-acute, (5–)6–9
× (0.8–)1–1.5 mm, softly to firmly coriaceous, flat or adaxially convex, with obscure to somewhat prominent vein
above, abaxially flat to slightly concave, with narrowly to broadly prominent vein, with flat to slightly revolute,
regularly, sometimes sparsely, denticulate margins. Leaf bases with prominently decurrent margins and vein. Upper
leaf epidermis cells ± isodiametric, thick-walled. Sporangia 1.5–2 mm wide.
Distribution:—Andes of Venezuela to southern Colombia, southern Ecuador.
Habitat:—Terrestrial, on wet road banks, land slides and rocky soil in subpáramo, elev. 3000–3400 m.
Notes:—Closely related to Phlegmariurus urbanii, from which it deviates by its larger proportions, fewer leaf
ranks, and the usually recurved but not sigmoid leaves. Resembling Phlegmariurus reflexus, but more robust, and with
more coriaceous leaves.
Phlegmariurus firmus, P. cleefianus, and P. urbanii are distinct from other members of the P. reflexus group due to
thick-walled, nearly isodiametric upper epidermal cells of the leaves. Phlegmariurus cleefianus differs from P. firmus
being nearly twice as large in most parts. Specimens examined:—Antioquia: And. Centr. Antioquia, 2500–3000 m,
Mayor 57 (S).
Mun. Sonsón, vereda Chaverras, slopes of quebrada El Padre Sanchéz, 2850–2900 m, Callejas & Roldán 10623
(AAU). Caldas: Nevado del Ruiz, Road Manizales–Hotel Termales, 3080 m, Santa & Hoyos 679 (AAU, COL, HUA,
QCA). Cauca: Cord. Central, Macizo Colombiano, E slope, Parque Nacional Puracé, ascent of S. José de Isnos, 2750
m, Cleef 5002 (AAU). Road to Parque Nacional Puracé, 3250 m, Santa & Hoyos 721 (AAU). Road Totoró–Inza, W
slope of Cord. Central, ascent to Páramo de Las Delicias, 3030 m, Santa et al. 972 (AAU). Road Totoró–Inza, E slope
of Cord. Central, descent from Páramo de Las Delicias, 3120 m, Santa et al. 980 (AAU). Mun. Puracé, ebtrance to
Pilimbalá–Laguna de San Rafael, 3200 m, Santa et al. 1002 (AAU). Macizo Colombiano, Valle de Las Papas, trail San
Sebastián–Valencia, N facing slope, 3000 m, Barclay & Juajibioy 5723 (GH, MO). Macizo Colombiano, Páramo de
Las Papas, surroundings of Laguna de La Magdalena, 3530 m, Idrobo et al 3351 (COL, NY). Páramo de Puracé, 8 km
E of Puracé, 25 km E of Popoyan, 3000–3300 m, Tryon & Tryon 5973 (COL, GH, NY, P, S, UC, US). Cundinamarca:
La Peña, Bogotá, 2900 m, Lindig 1521 (B, K, P). Huila: La Plata, 3000 m, Sneidern 2201 (S) Tolima: Road Mariquita–
Manizales, »Alto de Letras«, 3300 m, Santa & Escobar 856 (AAU, COL, HUA). Road Mariquita–Manizales, »La
Libia«, 3000–3200 m, Santa & Escobar 854 (AAU, QCA). E slope of Páramo de Letras, 3380 m, Cleef & Florschütz
5971 (AAU).
Phlegmariurus silverstonei B.Øllgaard (2016b: 269).—Fig. 25.
Type:—COLOMBIA. Chocó: Mun. San José del Palmar, Cerro del Torrá, filo de cumbre, vegetación abierta con arbustos, 2700–2800 m,
21 August 1988, Silverstone-Sopkin et al. 4652 (holotype CUVC, isotype AAU).
Description:—Plants ascending to erect, mature plants forming small clumps without basal, creeping and rooting
shoots, to 20 cm tall, to 3 times dichotomous. Shoots homophyllous, almost equally thick throughout, 8–13 mm in
diameter including leaves. Stems excluding leaves 3–4 mm thick at the base, sometimes slightly tapering upward,
completely concealed by leaves. Leaves uniform throughout, or slightly smaller upward, densely crowded, borne in
more or less regular, often oblique or obscure, alternating whorls of 8–10, these 0.5–1 mm apart, forming 16–20 often
obscure longitudinal ranks, subsigmoid, patent-ascending to imbricate from a short nearly perpendicular base, the
apex slightly falcate, linear-subulate, widest just above the base, (5–)6–8 × 0.6–1 mm, evenly tapering, with a broad,
prominent veinal ridge abaxially, adaxially flattened, without a basal swelling abaxially, not decurrent, green, with
smooth margins, the apical 1–1.5 mm minutely omnilaterally spiny, upper epidermal cells relatively thick-walled,
isodiametric or slightly longer than wide. Sporangia ca. 1.5 mm wide.
Distribution:—Known only from the type collection.
Habitat:—As indicated by the type label.
Notes:—This species shares the high number of leaves in leaf whorls with P. urbanii, and the pattern of upper
epidermal cells on the leaves, although slightly less pronounced in the present species .

FIGURE 25. A. Phlegmariurus silverstonei (Silverstone-Sopkin 4652, AAU, CUVC), growth habit. A1. Close-up of terminal sporangiate
division, with some leaves removed.
Phlegmariurus urbanii (Herter) B.Øllgaard (2012b: 20).—Fig. 10C.
Lycopodium urbanii Herter (1909a: 47).—Urostachys urbanii (Herter) Nessel (1939: 110).—Huperzia urbanii (Herter)
Holub (1985: 77). Type:—ECUADOR: Tungurahua: “inter saxa vulcanica”, elev. 10 000 ped., Spruce 5429 (P or K to
be chosen as lectotype; isotypes AWH, E, C, G, GH, NY, OXF).
Illustration: Øllgaard (1988: Fig. 2C).
Description:—Plants erect or erect from a decumbent base, small to very long, usually to 30 cm tall, seen to 100 cm
long, sparsely branched, to 4 times dichotomous. Shoots homophyllous, equally thick throughout, 6–10(–13) mm
in diameter including leaves. Stems excluding leaves 2–5(–8) mm thick at the base, sometimes tapering to 2–3 mm
thick, almost completely concealed by leaves, densely ridged by decurrent leaf bases, sporangiate from 6–30 cm above
the base and upward. Leaves uniform throughout, densely crowded, borne in alternating, more or less regular, often
oblique, whorls of (10–)11–14(–16), these 0.7–2(–3) mm apart, forming (20–)22–28(–32) obscure longitudinal ranks,
loosely imbricate-ascending to sharply reflexed, usually distinctly sigmoid, linear-subulate, often almost parallel-sided
to near the apex, short-acute, 3.5–4.5(–5) × 0.7–1 mm, thick and coriaceous, usually lustrous, adaxially convex with
obscure to widely prominent vein, abaxially with longitudinal grooves on both sides of the wide, strongly prominent
vein, with slightly revolute, sparsely denticulate to densely short and pale-ciliolate margins. Leaf base with prominently
decurrent vein and margins. Upper leaf epidermis cells ± isodiametric, thick-walled. Sporangia 0.8–1.2 mm wide.
Distribution:—Southern Colombia, Ecuador, Peru.
Habitat:—Terrestrial on land slides, road banks and other open, moist habitats in upper montane forest and
subpáramo, elev. 2300–3800 m.
Notes:—Phlegmariurus urbanii is a close relative of P. firmus, but has smaller, sigmoid leaves borne in a greater
number of orthostichies. Both species have short, thick-walled upper epidermal cells on the leaves, in contrast to the
usually thin-walled, elongate epidermal cells in P. reflexus and its relatives. Phlegmariurus silverstonei resembles this
species but is larger in all parts.
Specimens examined:—Cauca: El Tambo, carretera al mar por el veinte de Julio, flanco W de la cordillera
occidental, 2300 m, Santa et al 990 (AAU, COL, HUA, MEDEL). Nariño: Pasto, between El Encano and Santiago,
3100 m, Santa & Buitrago 934 (AAU, COL, HUA). S side of Volcán Dona Juana, 3000–3600 m, Ewan 16626 (US).
Putumayo: Road San Francisco–Mocoa, 2300 m, Bristol 408 (GH).
FIGURE 26. A. Phlegmariurus ascendens (Ecuador, Øllgaard & Balslev 8600, AAU). A1. Close-up of terminal sporangiate division.
Del K. Tind.

Group of Phlegmariurus tetragonus
This group includes two closely related terrestrial species, that share the feature of densely fimbriate leaf margins.
They are undoubtedly close to the following high Andean groups of Phlegmariurus crassus.
Phlegmariurus attenuatus (Spring) B.Øllgaard (2012b: 12).—Fig. 27B.
Lycopodium attenuatum Spring (1849: 8).—Huperzia attenuata (Spring) Trevisan (1874: 249).—Urostachys attenuatus (Spring) Nessel
(1939: 101). Type:—ECUADOR. Pichincha: “in declivitate montis Pichincha”, Hartweg 1470 (lectotype US, designated by Lellinger
(1977: 717), isotypes BM, GL, K, NY, P).
Lycopodium tobarii Sodiro (1893: 565).—Urostachys tobarii (Sodiro) Nessel (1939: 101). Type:—ECUADOR. Pichincha: “pasq. m.
Pichincha”, 3600 m, May 1874, Sodiro s.n. (lectotype QPLS, designated by Øllgaard 1988: 61).
Illustrations: Lellinger (1989: Fig. 15). Øllgaard (1988: Fig. 11D).
Description:—Plants ascending to erect, forming small clumps with erect shoots in the center, and more or less
prostrate-ascending, basal shoots, to 20 cm tall, or to 30 cm long, to 4(–5) times dichotomous. Shoots homophyllous to
slightly gradually heterophyllous, terete to hexagonal, equally thick throughout, 4–7 mm in diameter including leaves.
Stems excluding leaves 1.5–2 mm thick at the base, sometimes tapering to 1–1.5 mm in diameter upward, completely
concealed by leaves, sporangiate from few cm above the base, tinged with red–brick red. Leaves borne in alternating
whorls of 3–4, these 1–3 mm apart, forming 6 or 8 usually regular ranks, appressed to the stem throughout, in basal
divisions lanceolate and convex abaxially, upward gradually changing to narrowly triangular-ovate with acute–slightly
acuminate apex, abaxially strongly convex–subcarinate near the apex, with a prominent basal swelling (air sac), (4–)5–
6 × 1–1.5 mm wide, upward 1.5–2.2 mm wide, with fimbriate margins, tinged with red. Sporangia 1.5–2 mm wide.
Distribution:—Costa Rica, Andes from Colombia to Peru and Bolivia.
Habitat:—Terrestrial in exposed cushion vegetation in páramo, pioneer vegetation on land slides, road banks etc.
in the páramo zone, elev. 2800–4000 m.
Notes:—A confusing, heterogeneous assemblage of slender páramo species, including such a distinctive element
as Phlegmariurus hypogaeus, have been referred to this species in herbaria and earlier treatments. The lectotype,
however, belongs to an easily recognizable species, characterized by fimbriate leaf margins. Phlegmariurus attenuatus
is a close relative of P. tetragonus, which see for discussion.
Specimens examined:—Tolima: Rosalita, near Páramo de Ruiz, 2800–3100 m, Pennell 2952 (F, NY). Vargas
Gaviria, Buitrago & Esquivel (2018) recorded this species from Páramo de Anaime.
Phlegmariurus tetragonus (Hooker & Greville) B.Øllgaard (2012b: 19).—Fig. 27A.
Lycopodium tetragonum Hooker & Greville (1829: t. 109).—Lycopodium tetragonum Hooker & Greville var. patulum Spring (1849:
12).—Huperzia tetragona (Hooker & Greville) Trevisan (1874: 248).—Urostachys tetragonus (Hooker & Greville) Nessel (1939:
135). Type:—ECUADOR. Pichincha: “Prope verticem montis Pichincha”, Jameson s.n. (holotype E or K; isotype BM).
Lycopodium catharticum Hooker (1838: 430–431).—Urostachys catharticus (Hooker) Nessel (1939: 135).—Lycopodium myrsinites
Lamarck var. minus Spring (1842: 29).—Syntypes:—ECUADOR: Asuay of the Equator, Turner s.n. (K n.v.); ECUADOR, [prov.
Cañar] from the mountains of Pillzhum, Jameson s.n. (K).
Description:—Plants ascending to erect, forming small clumps with erect shoots in the center, and usually prostrate
to ascending basal shoots, to 20(–30) cm tall, 3–5 times dichotomous. Shoots homophyllous, or gradually slightly
heterophyllous, quadrangular and equally thick throughout, 2.5–3.5(–4) mm in diameter including leaves. Stems
excluding leaves 1–2 mm thick, sporangiate nearly from the base. Leaves decussate throughout, imbricate, concealing
the stem, in basal divisions widely lanceolate to trianguar-ovate, abaxially rounded and evenly decurrent, upward
triangular, carinate at least in the upper half, somewhat decurrent or abaxially with a prominent basal swelling (air sac),
3.5–5(–7) × 2–4 mm, evenly tapering into acute–slightly acuminate apex, orange to bright red tinged at the margins or
throughout, with fimbriate margins. An occasional branch may be hexagonal or terete, with leaves borne in alternating
whorls of 3. Sporangia 1.3–2.5 mm wide.
FIGURE 27. A. Phlegmariurus tetragonus (Ecuador, Holm-Nielsen et al. 5001, Ecuador, AAU), growth habit. A1. Close-up of sporangiate
division. A2. Two sporangiate leaves.—B. Phlegmariurus attenuatus, (Ecuador, Bell 705, S), growth habit. B1. Close-up of sporangiate
division. B2. Sporophyll. Del. B. Johnsen.
Distribution:—Colombia to Peru (a doubtful record from Bolivia). Vargas Gaviria, Buitrago & Esquivel (2018)
recorded this species from Páramo de Anaime, Cajamarca.
Habitat:—Exposed cushion vegetation, pioneer vegetation on land slides, road banks etc. in the lower part of the
páramos, elev. 2800–3740 m.
Notes:—The red or reddish, quadrangular, erect shoots make this species an easily recognizable one. Incomplete
material of some epiphytic species with quadrangular constricted shoots, have been frequently misidentified as this
species. They are readily separated by the fimbriate leaf margins in Phlegmariurus tetragonus.
Phlegmariurus tetragonus is closely related to P. attenuatus. The occasional aberrant hexagonal or terete shoots in
some specimens of P. tetragonus are virtually indistinguishable from slender shoots of P. attenuatus. The two species
are also ecologically rather similar and are often found growing intermixed. There are many examples of species in

which the number of leaves in each whorl is variable in the same population. The difference of terete and quadrangular
shoots in P. attenuatus and P. tetragonus may be due to such simple and taxonomically overrated variation within
populations of one species. The strong visual effect produced by the change in phyllotaxis, may not reflect a great
genetic difference.
Specimens examined:—Caldas: Villamaría, road Manizales–Nevado del Ruiz, 3740 m, Santa et al. 955 (AAU,
COL, HUA). Road Manizales–Hotel Termales del Ruiz, 3000–3500 m, Forero et al. 521 (COL). 35 km NE of
Manizales, 3460 m, Madison 1315 (GH). Páramo del Ruiz, 25 km SE of Manizales, 3400 m, Tryon & Tryon 6146
(GH). Nevado del Ruíz, road Páramo de Letras–Termales del Ruíz, N slope, 3300 m, Cleef & Hart 2372 (AAU).
Cauca: Páramo de Paletará, 3000 m, Espinal 3300 (COL). Sotara, Popoyan, Hartweg 1467 (BR, K, P, US). Huila/
Cauca: Macizo Colombiano, páramo de las Papas, mountains and vicinity of Laguna La Magdalena, 3530 m, Idrobo
et al. 3285 (COL). Macizo Colombiano, Valle de las Papas, near Valencia, 2910 m, Idrobo et al. 3817 (US). Tolima:
Rosalito, near Páramo de Ruiz, 2800–3100 m, Pennell 2952 (F, p.p., GH, K, NY p.p., US).
Group of Phlegmariurus crassus
I here unite the species in the clades including Phlegmariurus saururus and P. crassus in the cladogram of Testo et al.
(2018). I suspect that with additional species included in a future molecular study the two clades may well become
united or subdivided in different ways. All species in this group are terrestrial and belong to the high Andean plant
communities.
Phlegmariurus ascendens (Nessel) B.Øllgaard (2012b: 12).—Fig. 26.
Urostachys ascendens Nessel (1940a: 161).—Huperzia ascendens (Nessel) Holub (1985: 70). Type:—ECUADOR: Pichincha en los
páramos, Ecuador, Bernard 1909 (holotype BONN [Herb. Nessel 119]).
Description:—Plants with elongate, to more than 20 cm long, horizontal, creeping, rooting, isotomously branching
supraterranean shoots, from which stiffly erect, well spaced, aerial shoot systems arise. Creeping shoots loosely
covered by green, often somewhat secund leaves, 8–11 mm in diameter including leaves. Aerial shoots to 22(–30) cm
tall, to 4 times dichotomous, homophyllous, terete, equally thick throughout, 7–12 mm in diameter including leaves,
usually slightly tapering upward. Stems of aerial shoots 2–3 mm thick at the base excluding leaves, upward tapering
to 1.5–2 mm, partly concealed by leaves, brownish to bright red, smooth, with long-decurrent ridges from leaf bases.
Leaves borne in alternating, irregular whorls of 4–6, these 1–2.5 mm apart, forming 8–12 fairly regular, longitudinal
ranks, arcuate-appressed throughout or gradually closely appressed upward, linear-lanceolate in basal, non-sporangiate
divisions, upward gradually lanceolate to broadly lanceolate, widest short above the leaf base, evenly tapering to
slightly long-acuminate, in basal divisions 6–8 x 1.5–1.8 mm, upward (4.5–)5–7 × 1.5–1.8(–2.2) mm, adaxially flat to
concave (dried), abaxially convex, rounded, with a wide, prominent, long decurrent basal swelling (air sac), somewhat
lustrous, usually with red apex and margins, with smooth to unevenly rugose, sclerified margins, amphistomatic.
Sporangia 1.8–2.2 mm wide.
Distribution:—Northern Ecuador, southermost Colombia.
Habitat:—Terrestrial in grass páramos, elev. 3370–4050 m.
Notes:—The erect shoots of this species are somewhat similar to Phlegmariurus capellae, but its long-creeping,
rooting shoots are not found in P. capellae. Also its leaves in the aerial shoots are usually shorter, especially in the basal
divisions.
Specimens examined:—Nariño: Mpio. de Pasto, Estribaciones cordillera Centro-Oriental, Páramo Los Frailes,
3300–3350 m, Ramírez 10918 (PSO). This collection is referred here with some doubt.
Phlegmariurus brevifolius (Greville & Hooker) B.Øllgaard (2012b: 12).—Fig. 29.
Lycopodium brevifolium Greville & Hooker (1832: 104).—Urostachys rufescens (Hooker) Nessel var. brevifolius (Greville & Hooker)
Nessel (1939: 103).—Urostachys brevifolius (Greville & Hooker) Herter (1949: 54).—Huperzia brevifolia (Greville & Hooker)
Holub (1985: 71). Type:—According to protologue: PERU: in Herb. Lambert, Ruíz and Pavon s. n. (holotype FI-Webb; AAU [photo
of FI], BONN [Herb. Nessel 202, fragment of G], G).
Description:—Plants erect or ascending, stiff and robust, forming small to large clumps, without basal creeping
and rooting shoots, often more than 25 cm tall, to 7 times dichotomous. Shoots homophyllous, almost equally thick
throughout, 7–9(–10) mm in diam. including leaves, or rarely tapering to 5 mm in diameter in terminal divisions. Stems
excluding leaves (3–)4–7 mm thick at the base, slightly tapering upward, usually concealed by leaf bases, sporangiate
from 7–25 cm above the base and upward. Leaves almost uniform throughout, densely crowded, or sometimes more
spaced in basal divisions, borne in regular, alternating whorls of 4–5, these 0.7–1.5 mm apart, forming 8–10 longitudinal
ranks, ascending or perpendicular to reflexed, with straight to upward curved apex, broadly ovate or triangular-ovate to
broadly suborbicular-cordate or triangular-cordate, acute to almost obtuse or mucronulate, 2–4 × 2–4(–5) mm, stiffly
coriaceous, green or red tinged, abaxially concave, with sharply prominent vein, with sharply prominent, somewhat
flattened, short-decurrent basal swelling (air sac), with slightly revolute, strongly sclerified, darker and somewhat
translucent, slightly erose margins. Sporangia 1.5–2.5 mm wide.
Distribution:—Costa Rica, Venezuela to Peru. Ramírez Padilla & Macias Pinto (2007) reported this species from
Dept. Cauca.
Vargas Gaviria, Buitrago & Esquivel (2018) recorded this species from Páramo de Anaime, Dept. Tolima.
Habitat:—Wet páramos , elev. 3200–4200 m.
Notes:—The total distribution range above is given with some reservation, as there is considerable regionally
characteristic variation in the material referred to this species. Populations from Colombia, Southern Ecuador and Peru
are slightly different and usually recognizable, perhaps worth taxonomic recognition.
Phlegmariurus brevifolius differs from P. rufescens, P. serpentiformis, P. lignosus, and P. sellifolius by its larger
size, thicker stems, stiff and almost prickly leaves, and the leaves usually wider than long. Phlegmariurus idroboi is
somewhat similar but nearly twice as large.
Specimens examined:—Boyacá: Mun. Tutasá, high massif N of Belén, SE slopes of Cerro Pan de Azucar, 4100
m, Cleef 9810, 9815, 9816 (AAU). Páramo de la Rusia, NW–N of Duitama, Serranía Peña Negra, Hoya de la Laguna
Agua Clara, 3800 m, Cleef 7065 (AAU), ibid. 3950 m, Cleef 7435 (U). Páramos NW of Belén, headwaters of Quebrada
Minas, Hoya SE, 3910 m, Cleef 9716 (AAU). Mun. Socotá, Páramo Pisba, Peña Negra-lagoon Choro Negro, 3500 m,
Stancik & Medina 2341 (AAU). Cundinamarca: Páramo de Gutiérrez, 3290 m, Ariste-Joseph s. n. (BM, C). Cord.
Oriental S of Usme, Páramo de Chisacá, 4020 m, Barclay & Juajibioy 6212 (GH). Laguna de Chisacá, 3900–4200 m,
Bischler 1510 (BM). Valle: Cord. Central, vertient occidental, cabeceras de los ríos Tuluá y Bugalagrande, Páramo de
Las Vegas, 3600–3800 m, Cuatrecasas 20316 (F).
Phlegmariurus capellae (Herter) B.Øllgaard (2012b: 13).—Fig. 28.
Urostachys capellae Herter (1954: 114–115).—Lycopodium capellae (Herter) Morton (1964: 72).—Huperzia capellae (Herter) Holub
(1985: 71). Type:—ECUADOR: Napo [as Imbabura]: E of Volcán de Cayambe, along trail between Río Boquerón and Río Arturo,
11000 ft., Drew E 314 (lectotype US, designated by Rolleri (1981: 78), isolectotype, MSC).
Illustrations: Øllgaard (1988, Fig. 8B).
Description:—Plants ascending to erect, sparsely to densely branched, without prostrate-ascending, rejuvenating

shoots, to ca. 35 cm tall, to 6 times dichotomous. Shoots homophyllous to gradually slightly heterophyllous, 12–20
mm in diameter at the base including leaves, usually tapering to 5–10 mm in diameter, sporangiate from 8–20 cm
above the base and upward. Stems excluding leaves 2–3.5(–4) mm thick at the base, tapering to 1.5–2 mm thick,
often brown to reddish brown, terete to prominently ridged by decurrent leaf bases. Leaves of basal divisions (and
shaded shoots) borne in irregular, rather distant, alternating whorls of (4–)5–6, these 2–6 mm apart, forming (8–)10–12
longitudinal ranks, usually patent to ascending, not covering the stem, linear-lanceolate, 7–11 × 1.5–2 mm, adaxially
flat, lustrous, abaxially flat to slightly convex, dull to lustrous green, usually with long decurrent leaf bases. Leaves
of upper and terminal divisions gradually shorter, closer, and more appressed, borne in alternating whorls of 4–5(–6),
these 1–3 mm apart, forming 8–10(–12) longitudinal ranks, ascending to arcuate-appressed, straight to strongly upward
or unilaterally curved or twisted, lanceolate to widely lanceolate, often slightly cuspidate above the middle, (5–)6–8
× 1.5–2 mm, adaxially concave (dried) or flat to shallowly rounded in life, abaxially convex, rounded and irregularly
wrinkled (dried), with slightly prominent, short to long, narrowly decurrent basal swelling (air sac), amphistomatic,
with smooth to densely irregularly rugulate, slightly to distinctly, translucently sclerified margins, shining to dull
green. Sporangia ca. 2 mm wide.
FIGURE 28. A. Phlegmariurus capellae (Cuatrecasas 23176, S), growth habit. A1. Close-up of sporangiate division.
Distribution:—Andes from Venezuela to Peru.

Habitat:—Terrestrial in grass páramos, usually growing in partial shade among grasses, elev. 3200–3940 m.
Notes:—See discussion under Phlegmariurus crucis-australis.
Specimens examined:—Antioquia: Urrao, road to páramo de Frontino, 3320 m, Santa 1176 (AAU, HUA).
Urrao, road to páramo de Frontino, 3250 m, Santa 1183 (AAU). Urrao, páramo de Frontino, Alto de Campanas, 3820
m, Santa 1195, 1196 (AAU). Caldas: Nevado del Ruiz, Road Manizales–Nevado del Ruiz, 3640–3940 m, Santa &
Hoyos 684 (AAU, COL, HUA, MEDEL), 683, 693 (AAU, QCA). Cabeceras del río Otún, Laguna de Taburetes (arriba
de El Bosque), 3580 m, Cuatrecasas 23176 (F, S). Cauca: Road Totoró–Inza, páramo de Las Delicias, 3200 m, Santa
et al. 977 (AAU, COL, HUA, MEDEL). Cundinamarca: Río Colorado, 5 kms S de la Laguna de Chisacá, 3600 m,
Bischler 2373 (G). Nariño: Mun. Pasto, Volcán Galeras, 3600 m, Santa & Buitrago 942 (AAU, HUA). Putumayo:
Páramo del Tambillo. NE del valle de Sibundoy, 2700–2800 m, Schultes 3115 (GH, photo AAU). Risaralda: Cord.
Central, headwaters of Río Otún, Laguna de Taburetes (above el Bosque), 3580 m, Cuatrecasas 23176 (S, US).
Vargas Gaviria, Buitrago & Esquivel (2018) recorded this species from Páramo de Anaime, Dept. Tolima.
FIGURE 29. A. Phlegmariurus brevifolius (Cuatrecasas 20316, F), growth habit. A1. Close-up of sporangiate division.
Phlegmariurus catacachiensis (Nessel) B.Øllgaard (2012b: 13).—Fig. 30.
Urostachys catacachiensis Nessel (1940a: 162).—Huperzia catacachiensis (Nessel) B.Øllgaard (1987: 169). Type:—COLOMBIA:
“Ecuador, Catacachi, auf Grashalden” [probably a mislabelled Colombian plant], Bernard s.n. (holotype BONN [Herb. Nessel
181]).
Description:—Plants with age forming cespitose tufts, sometimes slightly heteroblastic with the heteroblastic shoots
ascending and not rooting, sometimes to 20 cm wide, 5–20 cm tall, at least to 6 times dichotomous. Aerial shoots
3–5(–8) mm in diameter including leaves, usually uniform throughout or slightly narrower upward. Aerial stems
1.5–2(–2.5) mm thick (dried) excluding leaves near the base, upward narrower, ca. 1–1.5 mm thick, usually red
and somewhat contrasting the leaves, stem surface usually minutely scabrous to densely papillose. Leaves of aerial
shoots borne in often somewhat irregular alternating whorls of (3–)4–5, these 1–3 mm apart, forming (6–)8–10 often
indistinct longitudinal ranks, homophyllous or gradually reduced upward, ascending to closely appressed, straight
to upward curved (often patent in shaded divisions), (3–)4–6(–7) × 1–1.2(–1.5) mm, subulate to lanceolate in basal
divisions, abaxially usually strongly convex or with a prominent vein (dried), slightly decurrent or with a prominent
leaf base (air cavity), upward (2.5–)3–6(–7) × (0.8–)1–1.2(–1.5) mm, gradually narrowly to broadly lanceolate, with a
prominent leaf base (air cavity), usually smooth, slightly lustrous or sometimes pruinous, adaxially concave, the apex
short-acute to narrowly acuminate, often slightly tinged with red, leaf margins smooth or somewhat rugate (rarely
minutely denticulate), amphistomatic. Sporangiate leaves usually with a prominent, basal swelling (air sac) causing a
perpendicular appearance of the very leaf base. Sporangia ca. 1–1.2 mm wide.
Distribution:—Colombia and Andean Venezuela.
Habitat:—Subpáramo and páramo, bogs in páramo. Elev. 2900—4200 m.

FIGURE 30. A. Phlegmariurus catacachiensis (Sneidern 2200, S), growth habit. A1. Vegetative division. A2. Sporangiate division.
Notes:—Plants usually with tufted growth, rarely heteroblastic. Shoots usually with closely appressed leaves
contrasting the red-tinged stem.
Specimens examined:—Antioquia: Mun. Urrao, Jaiperá, Vereda El Chuscal, Páramo de Frontino, Llanogrande,
3000—3390 m, Callejas et al. 7680 (AAU). Mun. Urrao, Páramo de Frontino, Llano Grande to Campañas and La
Laguna, including La Mosca, 3500—3800 m, MacDougal et al. 4525 (MO). Mun. Urrao, sector El Quince-Páramo
de Frontino, 2900—3400 m, Roldán 175 (MO, UC). Mun. Urrao, Páramo de Frontino, 3370 m, Santa 1186 (AAU).
Boyacá: Alto de Arcabuco, Romero s.n. (UPTC). Caldas: Road Manizales-Nevado del Ruiz, 3940m, Santa & S.
Hoyos 692 (AAU, COL, HUA, MEDEL, QCA). Páramo del Quindio, Cordillera Central, 3700–4200 m, Pennell &
Hazen 9929 (S). Road on N side of Nevado del Ruiz, that follows 4200 m contour, near fork to to El Rosario, ca. 8
km E of of intersection with road to Manizales, Funk 8050 (COL). Cauca: Paramo de Puracé, 3450 m, von Sneidern
2200 (S). Puracé, road to Parque Nacional de Puracé, Laguna de San Rafael, 3350 m, Santa & S. Hoyos 722 (AAU).
Cundinamarca: Usme, Laguna de Chisacá, 3550 m, Santa & Buitrago 874, 876 (AAU, COL, HUA). Páramo de
Verjón, above Bogotá, Bro. Ariste Joseph A743 (US). Laguna de Chisacá and surroundings, 3900—4200 m, Bischler
1511 (COL). Macizo de Sumapaz, N side, Andabobos, 3760—3720 m, Cuatrecasas 27035 (US). Páramo de Sumapaz,
páramo y bosque alto-andino cerca de Lagunitas al S de San Juan, 3450 m, Cleef 8435 (U). Región del Sumapaz, 2
km from road to San Juan, 3960 m, Rangel 1658 (COL). Páramo de Palacio, km 19 along road, cabeceras Río Negro,
3375 m, Cleef CI 03987 (AAU). Distr. Especial de Bogota: Páramo de Chisacá, near Laguna Chisacá, hoya del Río
Magdalena, 3700--3800 m, Pinto & Hernández 486 (COL). Meta: Páramo de Sumapaz, Cerro Nevado del Sumapaz,
W slope, 4000 m, Cleef 1332 (AAU, U). Páramo de Sumapaz, Cerro Nevado del Sumapaz, Orilla pantanosa de la
lagunita de cal del vertiente W-NW del Cerro, 3615 m, Cleef 1292 (AAU, U). Risaralda: Road along limit between
Nevado del Ruíz and Nevado de Santa Isabel, 3970 m, Santa et al. 945 (AAU). Valle del Cauca: Páramo de Las Vegas,
3600–3800 m, Cuatrecasas 20333 (F, S, US). Cord. Central, W slope, headwaters of ríos Tuluá and Bugalagrande:
Páramo de Las Vegas, 3600–3800 m, Cuatrecasas 20315 (S).
FIGURE 31. A-A1. Phlegmariurus cocuyensis (Type, Barclay & Juajibioy 7333, MO), growth habit. A2. Close-up of sporangiate
division.
Phlegmariurus cocuyensis B.Øllgaard sp. nov.—Fig. 31.
Diagnosis: Phlegmariurus cocuyensis is closely related to P. crassus, but differs usually by its smaller (but somewhat overlapping) size
in all parts, the leaves abaxially strongly rugose by protruding blister-like epidermal cells, and leaf margins coarsely and irregularly
erose.
Type:—COLOMBIA: Boyacá: Cordillera Oriental, Sierra Nevada del Cocuy, Alto Ritacuva, 4525 m, 11 April 1959, Barclay & Juajibioy
7333 (holotype MO).
Description:—Plants erect from a prostrate to ascending base, forming loose to dense clumps with basal, prostrate-
ascending, rooting shoots (these often absent in collections), and erect, finger-like aerial shoots, at least to 30 cm tall,
at least to 7 times dichotomous. Erect shoots homophyllous or almost so, equally thick throughout, or slightly tapering
upward, 5–7(–10) mm in diameter including leaves, sporangiate from ca. 5 cm above the base. Stems excluding leaves
2.5–3(–3.5) mm thick at the base, sometimes tapering to 2.5 mm thick, usually completely concealed by leaves, except

sometimes in basal or shaded divisions. Leaves uniform throughout, or slightly shorter upward, borne in irregular,
alternating whorls (or low spirals) of 4–6, these 1–2 mm apart, forming 8–12 obscure to regular longitudinal ranks,
densely crowded, usually closely imbricate, straight or upward curved, linear-lanceolate to triangular-lanceolate,
or ovate-lanceolate, narrowly to short-acute, (2.5–)3–5(–7) × 1–1.5(–2) mm, upward sometimes reduced to 2.5–3
mm long, with a prominent, short and prominently to long and evenly decurrent basal swelling (air sac) causing a
perpendicular appearance of the very leaf base in sporangiate leaves, amphistomatic, adaxially concave to slightly
convex, with a slightly raised veinal ridge near the base, abaxially convex and rounded or with a prominent veinal
ridge, strongly rugose by protruding, blister-like, sclerified epidermal cells, with rugose or irregularly erose, usually
strongly sclerified margins, green to dark red, often strongly pruinous. Sporangia 1–1.5 mm wide.
FIGURE 32. A. Phlegmariurus hypogaeus (Ecuador: Holm-Nielsen et al. 5343, AAU), growth habit. A1. Close-up of sporangiate
aerial branch.—B. Phlegmariurus crassus var. crassus (Ecuador: Holm-Nielsen et al. 5927, AAU. B1. Two sporangiate leaves. Del. B.
Johnsen
Distribution:—Endemic to the Eastern Andes of Colombia, most collections from Nevado del Cocuy.
Habitat:—Páramo and superpáramo, 3400–4700 m. Gravelly and rocky moraine.
Notes:—This species resembles Phlegmariurus crassus, and differs generally in leaf size, characters of the leaf
margins and abaxial leaf epidermis. There is considerable variation in plant and leaf size, but not least the leaf margin
characters seem distinct.
Two collections are aberrant (Sierra Nevada del Cocuy, Alto Ritacuva, 4100 m, Barclay 7366 (GH, MO) and
Bischler 2756 (COL, G)) with irregular leaf position, irregularly twisted, triangular to broadly lanceolate leaves,
possibly due to external damage to the apical meristem.
Specimens examined:—Arauca: Sierra Nevada del Cocuy, headwaters of Quebrada El Playón, Patio Bolos,
2.5 km SW of Laguna de La Plaza, hoya S. José, 4350 m, Cleef 9076 (AAU, U). Boyacá: Sierra Nevada del Cocuy,
Quebrada Bocatoma, 4230–4260 m, Cleef & Florschütz 5622 p.p., 5839, 5889 (AAU, U). Sierra Nevada del Cocuy,
Quebrada Bocatoma, 4100 m, Cleef 8735 (U). Nevado del Cocuy, alto valle de Las Lagunillas, Cuatrecasas & García
Barriga 1476 (COL, F). Nevado del Cocuy, Ritacuba, near Guicán, Las Playitas, 3950 m, Bischler 2764 (COL, G).
Nevado del Cocuy, Mun. Guicán, Filo este sobre laguna Pintada, 4300–4700 m, Bischler 2824 (COL). Sierra Nevada
del Cocuy, Valle de los Corallitos, 4300 m, Grubb & Guymer P 16 (COL). Sierra Nevada del Cocuy, Quebrada
Bocatoma, 4100 m, Cleef 8735 (AAU, U). Sierra Nevada del Cocuy, Páramo El Cóncavo, 4380 m, Kieft & Becerra
135 (AAU, COL). Sierra Nevada del Cocuy, Páramo Cóncavo, 4335 m, Cleef 8527 (U). Mun. Güicán, Sierra Nevada
del Cocuy, Laguna Grande de los Verdes, 3900–4100 m, Rangel et al. 1523 (COL). Sierra Nevada de Chita, Hda.
Ritacuba, 4500 m, Seminaristas Eudistas s. n. (COL). Road Cocuy–Chita, 3930 m, Santa & Escobar 1117 (AAU,
HUA). Mun. Guicán, Sierra Nevada del Cocuy, road to Cubará, 4200 m, Santa & M. Escobar 1114 (AAU, HUA).
Sierra Nevada del Cocuy, Páramo Cóncavo, Cuchilla Puentepiedra, ca 2 km NE of Laguna Pintada, 4510 m, Cleef
5681 (AAU, U). Santander: Páramo del Almorzadero, ca. 32–34 km S of Chitagá, ca. 3400 m, Luteyn et al. 7663 p.p.
(AAU).
Phlegmariurus crassus (Willd.) B.Øllgaard (2012b: 13).—Fig. 32B.
Lycopodium crassum Willdenow (1810: 50).—Urostachys crassus (Willd.) Nessel (1939: 75).—Huperzia crassa (Willd.) Rothmaler
(1944: 60). Type:—ECUADOR: Antisana, Humboldt & Bonpland 2263 (holotype B-Willd. no. 19417; isotypes BM, P).
Lycopodium erythraeum Spring (1849: 7).—Huperzia erythraea (Spring) Trevisan (1874: 249).—Urostachys erythraeus (Spring) Nessel
(1939: 74). Type:—ECUADOR: Antisana, Hartweg 1472 (lectotype K, designated by Øllgaard (1988: 54) isotypes BM, G, GL, P).
Urostachys crassus (Willd.) Nessel var. herteri Nessel (1934: 180). Type:—See Øllgaard (1988: 55) for a comment. The lectotype should
probably be chosen among the specimens in BONN [Herb. Nessel 123], all of which belong to Phlegmariurus crassus.
Urostachys bonae-voluntatis Herter (1954: 112).—Lycopodium bonae-voluntatis (Herter) Morton (1964: 72). Type:—ECUADOR:
Mojanda, Mille 95 (holotype US).
Urostachys pilgerianus Nessel (1940: 161).—Huperzia pilgeriana (Nessel) Holub (1985: 75). Type:—PERU: Cordillere, 1901, Rautenstock
et Mann s.n. (holotype BONN [Herb. Nessel 124]). See Øllgaard (1994: 34) for discussion of the type.
Urostachys springii Nessel (1940: 161).—Huperzia springii (Nessel) Holub (1985: 77). Type:—ECUADOR: Ecuador, Jameson 103
(lectotype BONN [Herb. Nessel 125], designated by Øllgaard 1988: 54). The lectotype is different from specimens in G and OXF
with the same collection number.
Illustrations:—Øllgaard (1988: Fig. 8C-D).
Description:—Plants erect from a prostrate to ascending base, forming loose to dense clumps with basal, prostrate-
ascending, rooting shoots (these often absent in collections), and erect, finger-like aerial shoots, at least to 40 cm tall,
at least to 7 times dichotomous. Erect shoots homophyllous or almost so, equally thick throughout, or slightly tapering
upward, 7–14 mm in diameter including leaves, sporangiate from 5–20 cm above the base. Stems excluding leaves
(2–)3–5 mm thick at the base, sometimes tapering to 2–3 mm thick, usually completely concealed by leaves, except
in shaded divisions or individuals. Leaves uniform throughout, or slightly reduced upward, borne in often irregular,
alternating whorls of 4–7, these 1–3(–4) mm apart, forming 8–14 obscure or oblique to regular longitudinal ranks,
densely crowded, usually closely imbricate, straight (or slightly upward curved in shaded divisions), linear-lanceolate
to lanceolate, or triangular-lanceolate, narrowly to broadly acute, (6–)7–11 × (1.5–)2–2.5 mm, upward sometimes
reduced to 6–8 mm long with a prominent, short to long decurrent basal swelling (air sac) causing a bulging or
perpendicular appearance of the very leaf base in sporangiate leaves, amphistomatic, adaxially concave to slightly
convex, with a raised veinal ridge near the base, abaxially convex and rounded or with a prominent veinal ridge,
slightly to strongly rugose by protruding, blister-like epidermal cells, with smooth to slightly rugose, usually sclerified
margins, green to brick red or dark red, usually strongly pruinous. Sporangia 1.5–2.5 mm wide.
Distribution:—Costa Rica to the Andes of Venezuela south to Bolivia.
Habitat:—Páramos, 3700–4600 m.
Notes:—Specimens from Mexico, Guatemala, and Hispaniola commonly referred this name apparently are a
different species.
The variation in this species is complex, and the material described above as Phlegmariurus cocuyensis may be
considered an extreme part of the species complex. The variety P. crassus var. manus-diaboli (B.Øllg.) B.Øllgaard
(2012b) is common in the páramos of northernmost Ecuador may be expected in the south of Colombia.
Specimens examined:—Caldas: Nevado del Ruíz, superpáramo, arenales 3 km al SW del Refugio, cabeceras Q.
Las Nereidas, 4400 m, Cleef 2458 (AAU). N side of Nevado del Ruíz, Just E of Río Azufradon road that crosses N
side of Volcano, 4000 m, Funk 8058 (COL). Páramo del Nevado del Ruíz, below Termales, 4200–4500 m, Humbert
et al. 27047 (COL, US). Páramo de Ruíz, 4300 m, Triana 696 (COL). Nevado del Ruíz, Trujillo 5562 (COL, HUA).
Manizales, Nevado del Ruíz, 4500 m, Yepes-Agredo 765 (COL). Páramo del Ruíz, higher slopes, 4200–4630 m, Barclay
5243 (COL). Cordillera Central, W slope, páramos del Nevado del Ruíz, 3900–4200 m, Cuatrecasas 9274 (COL,
F). Road Manizales–Nevado del Ruiz, 3760–4130 m, Santa & Hoyos 686, 695, 696 (AAU, COL, HUA, MEDEL,
QCA). Mun. Villamaría, road Manizales–Nevado del Ruiz, 3740 m, Santa & al. 956 (AAU). Páramo del Ruiz, Llano
5760 (HUA). Circo intermedio del Ruiz, 4000 m, Llano, 64 (US). Cord. Central, Páramo de Quindio, 3700–4400
m, 13 Aug 1922, Pennell 9801 (US). Páramo de Quindío, Cord. Central, 3700–4200 m, Pennell & Hazen 9928 (S,
US). Cordillera Central, W slope, páramos del Nevado del Ruíz, 3900–4200 m, Cuatrecasas 9274 (COL). Cauca:
Cordillera Central, W slope, Headwaters of Río Palo, high páramos between Quebrada de San Paulino and Quebrada
del López, Lagunillas de Las Casitas, 3700 m, Cuatrecasas 19085A (F, GH). Magdalena: Sierra Nevada de Santa
Marta, alrededores de cabeceras de río Sevilla, 3360–3570 m., Barclay & Juajibioy 6549 (GH, MO). Meta: Páramo de
Sumapaz, Cerro Nevado del Sumapaz del flanco N, 4170 m, Cleef 8153 (AAU). Nariño: Mun. Pasto, Volcán Galeras,
3950 m, Pinto et al. 1824 (COL). Mun. Pasto, Volcán Galeras, near TV antennas, 3900 m, Santa & L. F. Buitrago 937,
938 (AAU, HUA). Volcán de Chiles, 2560–3350 m, Ewan 16095 (NO). Risaralda: Road along Nevados del Cisne y
Santa Isabel, 3810 m, Santa et al 950 (AAU, COL, HUA, MEDEL). Parque Nacional de Los Nevados, Near Laguna
Otún, 3900–4450 m, Koops 238, CGK238 (AAU). Santander: Páramo de Las Vegas, 3600–3800 m, Cuatrecasas
20339 (S). Tolima: Páramo de Ruiz, 3800–4300 m, Pennell 3042 (K, MO, US). Nevado del Ruiz Massif, ESE of
Manizales, 5000 m, Archibald 4014 (E).
FIGURE 33. A. Phlegmariurus crucis-australis (Cuatrecasas 18988, S), growth habit. A1. Close-up of sporangiate division.
Phlegmariurus crucis-australis (Herter) B.Øllgaard (2012b: 14).—Fig. 33.
Urostachys crucis-australis Herter (1954: 119).—Lycopodium crucis-australis (Herter) C. Morton (1964: 72). Type:—COLOMBIA:
Cauca, cordillera central, vertiente occidental, cabeceras del río Palo, Alto de Las Casitas, 3800 m, Cuatrecasas 18988 (holotype
US-1854751; Isotypes A, F, GH, S).
Description:—Terrestrial, erect from a prostrate to ascending base, forming loose to dense clumps, at least to 30
cm tall, at least to 6 times dichotomous. Shoots homophyllous or almost so, equally thick throughout, or slightly
tapering upward, 8–15 mm in diameter including leaves, upward tapering to 6–10 mm sporangiate from 10–20 cm
above the base and upward. Stems excluding leaves ca. 3 mm thick at the base, sometimes tapering to 2 mm thick,
the stem surface smooth, usually partly concealed by leaves. Leaves uniform throughout, or slightly reduced upward,
irregularly arranged, spirally or in irregular alternating whorls of 3–4, these usually 2–3(–4) mm apart, forming 6–8
obscure longitudinal ranks, patent-ascending to imbricate, usually upward curved, linear-lanceolate to lanceolate, 6–8
× 1.5–2 mm, upward sometimes reduced to 6 mm long, with a long- and narrowly decurrent leaf base, amphistomatic,
adaxially concave to nearly flat, with a slightly raised veinal ridge near the base, abaxially convex and rounded with
smooth to minutely rugose margins, green, somewhat lustrous. Sporangia ca. 1.5 mm wide.
Distribution:—Endemic.
Habitat:—Páramo to superpáramo, 3800–4000 m.
Notes:—The material studied resembles Phlegmariurus capellae and P. cumingii, and in some respects appears
intermediate. It deviates by being more ramified from the base than the former, and by irregular leaf position and
orientation.
Specimens examined:—Meta: Páramo de Sumapaz, Cerro Nevado del Sumapaz, W slope, 4000 m, Cleef 1332
(U).
Phlegmariurus cruentus (Spring) B.Øllgaard (2012b: 14).—Fig. 34-37.
Lycopodium cruentum Spring (1849: 34).—Urostachys cruentus (Spring) Nessel (1939: 27).—Huperzia cruenta (Spring) Rothmaler
(1944: 59). Type:—COLOMBIA(?): “Nova Granada”, Purdie s.n. (lectotype K, designated by Øllgaard 1989: 40).
The type of Urostachys erikssonii Nessel (1940: 163) appears to belong here (COLOMBIA: Monte Ruiz, 3500 m, Schmidtchen s.n.
(lectotype BONN [Herb. Nessel 191], designated by Øllgaard 1989: 90). However the quality of the specimen is insufficient for a
definite decision.
Distribution:—Colombia (?), Venezuela (?). Vargas Gaviria, Buitrago & Esquivel (2018) recorded this species from
Páramo de Anaime, Dept. Tolima. Ramírez Padilla & Macias Pinto (2007) reported this species from Dept. Cauca.
Habitat:—Presumably restricted to paramos above 3000 m.
Notes:—The name Phlegmariurus cruentus is applied here to a highly diverse assembly of collections from
páramos and humid habitats above the forest limit. The variation appears to be associated with variation in substrate
quality (solid soil, rocks, boggy or peaty ground) affecting growth habit (homoblastic to heteroblastic) and with
exposure and light conditions. In addition, interbreeding is suspected to be fairly unlimited within the group, reflecting
the relative youth of this group in the high Andes. Thus much of the morphological variation seems to be individual
rather than based on breeding barriers between what might be considered separate taxa.
The type of Lycopodium cruentum Spring appears to be a young plant growing on a solid substrate, strongly
heteroblastic, and with low, slender aerial shoots. Three varieties are recognized:
Key to the varieties of Phlegmariurus cruentus
1. Plants predominantly heteroblastic.....................................................................................................................................................2

– Plants predominantly homoblastic .....................................................................................................................................................3
2. Prostrate shoots 6–8 mm in diameter including leaves, aerial shoots usually 3–5 mm in diameter including leaves, sporangiate
leaves 3–5 mm long, abaxially convex to subcarinate, usually appressed ...................................................................... var. cruentus
– Prostrate shoots 6–12 mm in diameter including leaves, aerial shoots 4–10 mm in diameter including leaves, sporangiate leaves
4–7 mm long, ascending to appressed, abaxially rounded or with slightly prominent vein .........................................var. illimitatus
3. Shoots usually irregularly branched, often with long, irregular intervals; leaves of sporangiate divisions usually irregularly twisted,
5–8 mm long ......................................................................................................................................................................... var. exilis
– Shoots usually branched with regular intervals, leaves of sporangiate divisions arcuate-appressed 3–5 mm long, usually straight,
not twisted ................................................................................................................................................................. var. innocentium

FIGURE 34. A. Phlegmariurus cruentus var. cruentus (Cleef 4375, U), growth habit. A1. Close-up of terminal sporangiate division.
Phlegmariurus cruentus (Spring) B.Øllgaard var. cruentus—Fig. 34.
Description:—Plants slightly to strongly heteroblastic, with age often forming cespitose tufts, to 30 cm tall, to at least
6 times dichotomous, young plants growing on solid substrate with prostrate rooting shoots bearing erect aerial shoots,
the prostrate shoots often missing in collections, older plants growing in soft substrates usually slightly heteroblastic
with ascending and usually not rooting basal peripheral shoots. Prostrate shoots 6–8 mm wide including leaves, with
leaves upward secund. Aerial shoots 3–5(–6) mm in diameter including leaves, usually uniform throughout or slightly
narrower upward. Aerial stems 1–2 mm thick excluding leaves near the base, upward narrower, ca. 1–1.5 mm thick,
pale greenish or often reddish tinged. Leaves of aerial shoots borne in regular to slightly irregular alternating whorls of
3–4, these 1–1.5 mm apart, forming 6–8 often distinct longitudinal ranks, homophyllous or gradually reduced upward,
ascending or usually appressed, straight from a nearly perpendicular base to upward curved, 4–6(–7) × 0.8–1.2 mm,
subulate to lanceolate in basal divisions, upward 3–5 × 1–1.2 mm, gradually narrowly to broadly lanceolate, abaxially
convex or with a prominent vein or subcarinate (dried) and decurrent, usually smooth and slightly lustrous, sometimes
whitish pruinous, adaxially concave, the apex acute to narrowly acuminate, often tinged with red, leaf margins smooth
or somewhat. Sporangiate leaves usually with a slightly prominent, basal swelling (air sac) causing a perpendicular
appearance of the very leaf base. Sporangia ca. 1.5 mm wide.
Distribution:—Andes of Colombia, Venezuela.
Habitat:—Páramo and peat bogs, elev. (2800)3100–4200 m, mainly at low elevations in the range.
Specimens examined:—Boyacá: Páramo de Pisva, road Socha—La Punta km 69, between El Cardón and El
Cadillal, 3560 m, Cleef 4646 p.p. (U). Páramo de Pisva, road Socha–La Punta km 61.5, 6 km E of Los Pinos Alto
de Granados, 3570 m, Cleef 4375 (U). Páramo de Chita, headwaters of Río Casanare, hoya Laguna Chorros Blancos
ca. 2 km NE of Peñón Debisques, 3340 m, Cleef 9930 (AAU). Cundinamarca: Usme, Páramo de Sumapaz, 3500
m, Schneider 1012 (COL, S). Río Colorado, 5 km S of Laguna de Chisacá, 3600 m, Bischler 2374 (COL). Norte de
Santander: Páramo Almorzadero, 4200 m, Vareschi 4048 (VEN). Ibid. Vareschi 6046 (VEN). Tolima: ”Rosalito” near
Páramo de Ruíz, 2800—3100 m, Pennell 2958 (NY).
Phlegmariurus cruentus (Spring) B.Øllgaard var. exilis B.Øllgaard var. nov.—Fig. 35.
Diagnosis: This variety deviates from var. cruentus by lax, often tufted growth, and usually long slender and sparsely ramified aerial stems,
and irregularly alternating leaf whorls, and leaves often irregularly twisted.
Type:—COLOMBIA. Boyacá: Páramo de Guantiva, Ciénaga del Visitador, Carretera Sta. Rosita–Onza, 4 km NW de Sta. Rosita, 3290
m, Cleef 9860 (holotype AAU).
FIGURE 35. A. Phlegmariurus cruentus var. exilis (Cleef 9860, AAU), growth habit. A1. Close-up of basal segment. A2. Close-up of
terminal sporangiate segments.

Description:—Plants slightly to strongly heteroblastic, with age sometimes forming loose cespitose tufts, to 30 (–35)
cm tall, aerial shoots to 4(– 6) times dichotomous. Ascending or prostrate rooting shoots bearing erect aerial shoots, the
prostrate shoots often missing in collections, older plants growing in soft substrates usually slightly heteroblastic with
ascending rooting shoots. Prostrate shoots 6–10 (–12) mm wide including leaves, with leaves usually upward secund.
Aerial shoots 4–7(–12) mm in diameter including leaves, uniform throughout or slightly narrower upward. Aerial
stems 1–1.5(–2) mm thick excluding leaves near the base (dried), upward narrower, ca. (0.5–)1–1.5 mm thick, pale
greenish. Leaves of aerial shoots borne in often irregular alternating whorls of 3–5, these 1–2(–2.5) mm apart, forming
6–10 often indistinct longitudinal ranks, homophyllous or gradually reduced upward, ascending to appressed, often
patent in shaded divisions, straight to upward curved and often irregularly twisted, (4–)5–8 × 0.8–1.2 mm, subulate
to lanceolate in basal divisions, upward 4–7 × 1–1.2(–1.5) mm, gradually narrowly to broadly lanceolate, abaxially
convex and rounded or with a prominent vein (dried) and decurrent, usually smooth and slightly lustrous, sometimes
greyish pruinous, adaxially concave, the apex acute to narrowly acuminate, often tinged with red, leaf margins smooth
or slightly rugate. Sporangiate leaves usually with a slightly prominent, basal swelling (air sac) causing a perpendicular
appearance of the very leaf base. Sporangia 1–1.5 mm wide.
Habitat:—Moist to wet subpáramo and páramo, road banks, peat bogs, Sphagnum bogs. Elev. 3200–4000 m.
Specimens examined:—Boyacá: Nevado del Cocuy, Mun. Guicán, Corralitos, Salto de San Pablino, 4000 m,
Bischler 2956 (NY). Páramo de Guantiva, Ciénaga del Visitador, road km 4 NW of Sta. Rosita on road to–Onza,
3290 m, Cleef 9860 (AAU). Páramo de La Rusia, NW–N de Duitama, Hoya Laguna Negra, 1.5 km NNE of Laguna
Negra, Cleef 7507 (AAU). Páramo de La Rusia, NW-N de Duitama, 3490 m, Cleef 6832 (AAU). Sierra Nevada de
Cocuy, Valle de las Playas, 4200 m, Grubb & Guymer P. 43 (COL). Sierra Nevada del Cocuy, Alto de la Cueva, 3885
m, Cleef 9971 (AAU). Sierra Nevada del Cocuy, Páramo Cóncavo, valle del Río Cóncavo, 3770 m, Cleef 10009
(AAU). Cundinamarca: Páramo de Cruz Verde, ca. 10 km by road E of Bogotá, on road to Choachi, 3050–3200 m,
Duncan 1875 (UC). Páramo de Choachi, 3200–3300 m, Pennell 2240 (NY). Bogotá–Usme road, km 28–33, Páramo
de Chisacá, 3375–3450 m, Luteyn et al. 7750 (AAU, NY). Páramo de Chisacá, 3700–3800 m, Pinto & Hernandez
487 (COL). Laguna de Chisacá, 3650 m, Acosta-Arteaga 369 (AAU). Páramo de Cruz Verde, Turbera N of Laguna El
Verjón, 3455 m, Cleef 3200 (AAU). Páramo de Cruz Verde, road Bogotá–Choachí, km 15, 3355 m, Cleef 3060 (AAU,
U). Machetá, Páramo de Güina, 3200 m, Rangel et al. 4058 (MO). Dist. Especial de Bogotá, Páramo de Chisacá, lomas
cerca a la Laguna, Murillo & Fayad 34 (COL, F). Macizo de Bogotá, Páramo de Chisacá, around Laguna Chisacá,
3650–3700 m, Cuatrecasas & Jaramillo 25730 (COL, F, NY). Guadalupe, 5 km E of Bogotá, 3320 m, Little & little
9188 COL, F). Norte de Santander: Páramo de Romeral, 3800–4000? m, Killip & Smith 18662 (COL, S).
Phlegmariurus cruentus (Spring) B.Øllgaard var. illimitatus B.Øllgaard var. nov.—Fig. 36.
Diagnosis: This variety deviates from var. cruentus by an irregular growth habit, with often densely foliose basal divisions and short to
long, irregularly branching aerial divisions, and longer, patent to arching, regularly situated leaf whorls.
Type:—COLOMBIA: Páramo de La Rusia, NW–N de Duitama, 3490 m, Cleef 6832 (holotype AAU, isotype U).
Description:—Plants slightly to strongly heteroblastic, with age forming cespitose tufts, 5–28 cm tall, sometimes to
20 cm or more wide, at least to 6 times dichotomous, young plants usually with prostrate rooting shoots bearing erect
aerial shoots, the prostrate shoots often missing in collections, or plants from soft substrates slightly heteroblastic and
with ascending and not rooting peripheral shoots. Prostrate shoots 6–10 (–12) mm wide including leaves, with leaves
upward secund. Aerial shoots (3–)4–10(–15) mm in diameter including leaves, usually uniform throughout or slightly
narrower upward. Aerial stems (1–)1.5–2.5 mm thick excluding leaves near the base, upward narrower, ca. 1–2 mm
thick, pale stramineous to greenish. Leaves of aerial shoots borne in often somewhat irregular alternating whorls
of (3–)4–5, these 1–3 mm apart, forming (6–)8–10 often indistinct longitudinal ranks, homophyllous or gradually
reduced upward, ascending to appressed, straight to upward curved (often patent in shaded divisions and in dried
specimens), (4–)5–10 × 0.8–1.2(–2) mm, subulate to lanceolate in basal divisions, upward 3–7 × 1–1.2(–1.5) mm,
gradually narrowly to broadly lanceolate, abaxially rounded or with a prominent vein (dried) and decurrent, usually
smooth and slightly lustrous, sometimes whitish pruinous, adaxially concave, the apex acute to narrowly acuminate,
often slightly tinged with red, leaf margins smooth or somewhat rugate (rarely minutely denticulate). Sporangiate
leaves usually with a slightly prominent, basal swelling (air sac) causing a perpendicular appearance of the very leaf
base. Sporangia 1.5–2 mm wide.
FIGURE 36. A. Phlegmariurus cruentus var. illimitatus (Cleef 6832, AAU), growth habit. A1. Close-up of terminal sporangiate
divisions.
Habitat:—Subpáramo to superparamo, humid and humus-rich, peaty and boggy substrates, elev. 3100–4260 m.
Notes:—The remarks about the apparent limitless variation mentioned under Phlegmariurus cruentus above
applies not least to the present variety, and I suspect that future detailed field studies may reveal features that may
serve to separate entities, that so far remain obscure to the present author.
Specimens examined:—Boyacá: Páramo de Pisva, road Socha–La Punta km 69, between El Cardón and El
Cadillal, 3560 m, Cleef 4646 p.p. (U). Páramo de Pisva, road Socha–La Punta km 61.5, 6mkmE of Los Pinos Alto
de Granados, 3570 m, Cleef 4375 (U). Mpio. Duitama, páramo de La Rusia, 3540 m, Santa & Buitrago 887 (AAU).
Páramo de La Rusia, NW-N of Duitama, 3490 m, Cleef 6832 (AAU, U). Páramos al NW de Belén, cabeceras Quebrada
Minas, 3785 m, Cleef 1842 (AAU, U). Páramos al NW de Belén, cabeceras Quebrada Minas, 4050 m, Cleef 1939
(AAU, U). Road Sogamoso–Pajarito, headwaters of Río Cusiana, La Horqueta, subparamo 10 km NW of Vado Hondo,
3100 m, Cleef et al. 9251 (AAU, U). Vado Hondo, Siberia, páramo entre Peña de Arical y Alto de Mogotes, 1 km
al S de la Laguna Grande, 3320 m, Cleef 9260 (AAU, U). Sierra Nevada del Cocuy, Quebrada Bocatoma, 4260 m,
Cleef & Florschütz 5844 AAU, U). Cundinamarca: Mpio. de La Calera, páramo de Palacio, 3500–3800 m, Santa
1028 (AAU). Mpio. Zipaquirá, alto del Chamisero, 3200 m, Santa 1023 (AAU). Páramo de Chisacá, SE of Bogotá,
3350–3650 m, Schultes 20173 (COL). Mpio. de Usme, laguna de Chisacá, 3500 m, Santa & Buitrago 872 (AAU).
Usme, Páramo de Sumapaz, 3500 m, Schneider 1012 (S). Macizo de Sumapaz, near Lauguna Mayor, 3600 m, Uribe
6793 (COL). Páramo de Cruz Verde, road Bogotá–Choachí km 14, 3335 m, Cleef 2933 (AAU). Páramo de Palacio, km
19 de la carretera, Cabeceras Río Negro, 3370 m, Cleef 3976, 3987 (AAU). Páramo de Palacio, 1.5 km S of Lagunas

de Buitrago, 3560 m, Cleef 4121 (AAU). Meta: Páramo de Sumapaz, Hoys El Nevado, Lagunas El Sorbedero y El
Nevado, 3560 m, Cleef 1500 (AAU). Tolima: “Rosalito” near Páramo del Ruiz, 2800–3100 m, Pennell 2958 (COL,
NY).
Phlegmariurus cruentus var. innocentium (Herter) B.Øllg. stat. nov.—Fig. 37.
Urostachys innocentium Herter (1954: 127).—Lycopodium innocentium (Herter) C. Morton (1964: 72). Type:—COLOMBIA:
Cundinamarca, Páramo Gutiérrez, 3290 m, Bro. Ariste Joseph s.n. (holotype US 1342101)
FIGURE 37. A. Phlegmariurus cruentus var. innocentium (Jaramillo Mejia 2785, COL), growth habit. A1. Close-up of terminal
sporangiate division
Description:—Plants homoblastic or slightly heteroblastic, forming cespitose tufts, to ca. 30 cm tall, to ca. 20 cm or
wider, at least to 6 times dichotomous, heteroblastic shoots usually missing in collections, and when present ascending
and rarely or not rooting. Aerial shoots 5–8(–10) mm in diameter including leaves, uniform throughout or slightly
narrower upward. Aerial stems 1–1.5(–2) mm thick excluding leaves near the base, upward sometimes narrower, ca.
1–1.5 mm thick, pale greenish. Leaves of aerial shoots borne in often somewhat irregular alternating whorls of 4–(5),
these 1–2(–3) mm apart, forming 8(–10) often indistinct longitudinal ranks, homophyllous or slightly and gradually
reduced upward, patent to ascending in basal divisions, upward more arcuate- appressed, straight to upward curved
(often patent in shaded divisions and in dried specimens), 4–6 × 0.8–1.3 mm, subulate to lanceolate in basal divisions,
upward (2.5–)3–5 × 0.8–1.2(–1.5) mm, narrowly to broadly lanceolate, abaxially convex or with a prominent vein
(dried) and decurrent, usually smooth and slightly lustrous, sometimes greyish pruinous, adaxially concave, the
apex acute to narrowly acuminate, often tinged with red, leaf margins smooth or somewhat rugate (rarely minutely
denticulate). Sporangiate leaves usually with a slightly prominent, basal swelling (air sac) causing a perpendicular
appearance of the very leaf base. Sporangia 1–1.5 mm wide.
Habitat:—Boggy, marshy, poorly drained, mossy vegetation in páramo, elev. 3300–4050 m.
Notes:—This variety generally presents amply branched and elongate aerial branch systems with regular whorls
of rather short patent leaves.
Specimens examined:—Boyacá: Above Laguna de Tota, near Las Cintas, 3500 m, Jaramillo Mejía et al. 2785
(COL). Páramos al NW de Belén, cabeceras Quebrada Minas, 3910 m, Cleef 9728 (AAU, MO, U). Cundinamarca:
Páramo Palacios, Al E de La Calera, 3800 m, Hagemann & Leist 1301 (COL). Páramo de Palacio, km 19 along road,
headwaters of Río Negro, Cleef 3987. (AAU, U). Mun. Usme, laguna de Chisacá, 3550 m, Santa & Buitrago 874,
875 (AAU). Páramo entre Cogua y San Cayetano, cercanías de la Laguna Seca, Cabeceras Río Guandoque, 500 m
E de Laguna. Vallecito pantanoso, común, 3650 m, Cleef 6111 (AAU, U). Santander: Municipio Toná, Serranía de
Santurbán, road Berlín-Vetas, páramo de La Calaverita, 3600 m, Santa & Escobar 1097 (AAU).
Phlegmariurus cumingii (Nessel) B.Øllgaard (2012b: 14).—Fig. 38.
Urostachys cumingii Nessel (1934: 178).—Lycopodium cumingii (Nessel) Rolleri (1981: 92).—Huperzia cumingii (Nessel) Holub (1985:
72). Type:—ECUADOR: Quito, an bemoosten Felswänden, 1887, Cuming s n. (lectotype BONN [Herb. Nessel 51], designated by
Øllgaard 1988: 50).
Description:—Plants erect from a decumbent base, forming small to very large, dense clumps, with short basal,
prostrate-ascending often rooting shoots (these often absent in collections), to 35 cm tall, to at least 6 times dichotomous.
Shoots homophyllous and equally thick throughout, 6–15 mm in diameter including leaves near the base, or gradually
slightly heterophyllous and tapering to 4–8 mm in diameter upward, sporangiate from 5—15 cm above the base and
upward. Stems excluding leaves 1.5–3(–4) mm thick at the base, sometimes tapering upward to ca. 1–1.5 mm thick,
partly to completely concealed by the leaves. Leaves borne in irregular, alternating whorls of 5 or 6, these 1–2(–3) mm
apart, forming 10–12 indistinct longitudinal ranks, loosely to densely crowded, ascending to usually densely imbricate,
sometimes spreading in shaded shoots, almost straight to upward curved, linear-lanceolate to lanceolate, 5–7.5(–9) ×
1.2–1.5(–1.8) mm in basal divisions, upward usually reduced to 3.5–6 × 1–1.2 mm, adaxially concave or flat, with
slightly prominent vein, abaxially rounded or with a prominent veinal ridge, with slightly prominent, long decurrent
basal swelling (air sac), amphistomatic, with smooth to irregularly uneven, narrowly or not sclerified margins, usually
glossy green to yellowish or orangish green, sometimes with red or orange tinged margins and tips, rarely pruinous.
Sporangia 1.2–1.5 mm wide.
Distribution:—Ecuador and Colombia
Habitat:—Humid to wet grass páramos, 2900–4100 m, often growing in the shade of bunch grasses.
Notes:—Phlegmariurus cumingii usually forms densely branched, very large clumps of rather slender, green to
yellowish green slightly glossy shoots.
Specimens examined:—Cauca: Páramo de Buena Vista, Huila group, Central Cordillera, 3000–3600 m, Pittier
1113 (US). Cundinamarca: 4 km NW de Zipaquirá, páramo húmedo, Patano Redondo, 3100–3200 m, Tryon & Tryon
6062 (GH, NY, P, S, UC, US). Páramo de Guasca, 7 km E of Guasca, 3000–3100 m, Tryon & Tryon 5931 (COL, UC,
US). Valle: Cordillera Central, vertiente occidental; cabeceras de los ríos Tuluá y Bugalagrande, Páramo de Las Vegas,
3600–3800 m, Cuatrecasas 20290 (F, P, S, US). Province unknown: Mutis 3347 (US).
FIGURE 38. A. Phlegmariurus cumingii (Cuatrecasas 20290, F), growth habit. A1. Close-up of sporangiate division.
Phlegmariurus dianae (Herter) B.Øllgaard (2012b: 14).—Fig. 39.
Urostachys dianae Herter (1954: 116).—Lycopodium dianae (Herter) Morton (1964: 72).—Huperzia dianae (Herter) B.Øllgaard (1987:
169).—Lectotype.—COLOMBIA: Dept. Antioquia, Páramo of Morro Frontino, N of Urrao, elev. 11800 ft, 10 March 1944, E.
L. Core 388 (lectotype US-2290006, designted here, isolectotype US-2203759). First designated by C. Morton in handwriting,
indicating that Herter apparently forgot to annotate the specimens in US. Herter indicated a duplicate in Herb. Basel (not seen).
Description:—Plants erect or ascending, often forming large clumps, to at least 5 times dichotomous, without basal,
creeping and rooting shoots, sometimes more than 25 cm tall. Shoots homophyllous, almost equally thick throughout,
8–12 mm in diameter including leaves. Stems excluding leaves 2.5–4(–5) mm thick, almost completely concealed
by leaf bases, except in basal divisions, sporangiate from 5–15 cm above the base and upward, the stem surface
often papillose. Leaves almost uniform throughout, or gradually shorter and wider upward, usually densely crowded
throughout, borne in rather regular alternating whorls of 5–6, these 1–1.5 mm apart, forming 10–12 longitudinal
ranks, loosely imbricate or ascending to patent, sometimes somewhat sigmoid to recurved, ovate-lanceolate (in basal
divisions) to ovate, ovate-triangular or lanceolate-elliptic, acute or obtuse (or appearing obtuse due to incurved apex),
1.5–2 × 3–4(–5) mm, subcoriaceous to coriaceous, sometimes pruinous, sometimes slightly red tinged, abaxially
concave with prominent vein in basal part of lamina, to convex at the apex or sometimes convex nearly throughout, with
prominent to sharply prominent vein, short to long-decurrent basal swelling (air sac), margins flat to slightly involute
(rarely narrowly revolute), smooth or slightly to minutely rugose by individually protruding marginal, adaxially flat to
somewhat concave, or with prominent vein, hypostomatic. Sporangia 1.5–2 mm wide.
FIGURE 39. A. Phlegmariurus dianae (Type, US), growth habit. A1. Close-up of terminal sporangiate divisions.
Distribution:—Colombia. A local and variable species of the lower Páramos de Frontino.

Habitat:—Páramo, 2900–3550 m.
Notes:—Leaves of smaller and juvenile individuals often narrower (elliptic) than those of large and densely
sporangiate divisions. Closely related to the somewhat larger Phlegmariurus josesantae and often co-occurring with
that.
Specimens examined:—Antioquia: Mun. Urrao, sector El Quince-Páramo de Frontino, 2950–3400 m, Roldán
173, 174 (MO). Mun. Urrao, trail Finca La Quinche-Páramo de Frontino, 3200–3600 m, McPherson et al. 13128
(AAU). Mun. Urrao, Páramo de Frontino, Llano Grande, 3500 m, Roldán et al. 374 (MO). Mun. Urrao, Páramo
de Frontino, 3400 m, Londoño et al. 303 (HUA). Mun. Urrao, Páramo de Frontino, Llano Grande–Churrumblúm,
3320–3550 m, MacDougal et al. 4366 (MO, UC). Páramo de Frontino, 3660–3960 m, Carriker s.n. (US). Mun. Urrao,
Páramo de Frontino, 3450 m, Santa, J. 1177 (AAU), 1179 (AAU, QCA). Mun. Urrao, Páramo de Frontino, 3580 m,
Santa 1190 (AAU). Mun. Urrao, Páramo de Frontino, Inspección Jaiperá, Vereda El Chuscal, Llanogrande, 3400 m,
Callejas et al. 7631 (AAU).

Phlegmariurus eremorum (Rolleri) B.Øllgaard (2012b: 14).—Fig. 40.
Lycopodium eremorum Rolleri (1978: 141).—Huperzia eremorum (Rolleri) Holub (1985: 72). Type:—COLOMBIA. Valle del Cauca:
Cordillera Occidental, Los Farallones, entre La Torre y Alto del Buey, 3550–3400 m, Cuatrecasas 21907 (holotype US; isotypes F,
S).
FIGURE 40. A. Phlegmariurus eremorum (Cuatrecasas 17875, S), growth habit. A1. Close-up of sporangiate division.
Description:—Plants erect or ascending, forming small clumps, to 20–30 cm tall, usually 2–3 times dichotomous,
without basal, creeping and rooting shoots. Shoots almost homophyllous, almost equally thick throughout, 7–11 mm
in diameter including leaves. Stems excluding leaves 3–3.5 mm thick, almost completely concealed by leaf bases,
sporangiate from 10–15 cm above the base and upward. Leaves densely crowded throughout, borne in rather regular,
alternating whorls of 4(–5), these 1–1.5 mm apart, forming 8(–10) usually distinct longitudinal ranks. Leaves uniform
throughout or slightly shorter upward, ascending to arcuate-appressed, ovate and obtuse with upward-curved apex,
3.5–5 × 2–2.5 mm, hypostomatic, abaxially rounded or with slightly prominent vein, with narrowly decurrent base and
a small prominent air sac, adaxially concave, softly to firmly coriaceous, brightly red tinged, adaxially with a slightly
prominent veinal ridge, with usually narrowly sclerified, irregularly rugose margins. Sporangia ca. 1–1.4 mm wide.
Habitat:—
Notes:—Resembling Phlegmariurus rufescens and P. sellifolius, but larger and less ramified than these, and with
uniformly ascending-appressed, abaxially convex leaves.
Specimens examined:—Valle: Cordillera occidental, Los Farallones, crest of Cerro La Torre, 3750 m, Cuatrecasas
17875 (F, S, US, photo AAU).
Phlegmariurus hohenackeri (Herter) B.Øllgaard (2012b: 15).—Fig. 41.
Lycopodium hohenackeri Herter (1909a: 46).—Urostachys hohenackeri (Herter) Nessel (1939: 105).—Huperzia hohenackeri (Herter)
Holub (1985: 73). Type:—PERU: “in summis Cordillera jugis pr. Tabina”, July 1854, Lechler, pl. peruvianæ, ed. Hohenacker 2043
(B, BR, K n. v., NY fragment, P00559141, P00559140, S).
Illustrations: Øllgaard (1988: Fig. 13A, 13D).
Description:—Terrestrial, erect or ascending, forming small to large clumps, at least to 30 cm tall, at least to 3 times
dichotomous, without basal creeping and rooting shoots. Shoots homophyllous, equally thick throughout, 10–12 mm
in diameter including leaves. Stems excl. leaves 2–3 mm thick at the very base, upward increasing to ca. 4 mm thick,
usually completely concealed by leaves, sporangiate from about 10 cm above the base and upward. Leaves densely
crowded, borne in rather regular, alternating whorls of 4–5, these 1–1.5 mm apart, forming 8–10 usually regular
longitudinal ranks. Leaves of basalmost disions sublate to ovate, to 4 x 1.5—2 mm ascending to spreading, in upper
and terminal divisions broadly triangular-lanceolate to ovate-lanceolate, acute, 3–5 x 2.5–3 mm, with short prominent
leaf bases, loosely imbricate, softly to firmly coriaceous, red tinged, adaxially concave with a slightly prominent veinal
ridge, abaxially usually slightly concave at the very base but with a bulging air sac, with upward curved apex, with
usually strongly sclerified, irregularly rugose to minutely erose margins. Sporangia 1.5–2 mm wide.
Distribution:—Colombia, Ecuador, Peru. Vargas Gaviria, Buitrago & Esquivel (2018) recorded this species from
Páramo de Anaime, Dept. Tolima.
Habitat:—Páramos, 4200–4500 m.
Notes:—Phlegmariurus hohenackeri appears most closely related to P. brevifolius, with equally or even thicker
stems, but with larger, thinner, narrower, and usually uniformly ascending leaves. A variable species, or perhaps better
a species complex, poorly represented in collections.
Specimens examined:—Boyacá: Sierra Nevada El Cocuy, lado sur de La Plaza, arriba del acampamento, 4500
m, Hammen & González 1337 (COL). Habitat photo in AAU by Edgar Rincón Barón from same locality at 4200 m
elev.
Phlegmariurus hypogaeus (B.Øllg.) B.Øllgaard (2012b: 16).—Fig. 32A.
Huperzia hypogaea B.Øllgaard (1988: 58). Type:—ECUADOR. Carchi: … Øllgaard & Balslev 8517 (holotype AAU; isotype QCA).
Illustrations: Øllgaard (1988: Fig. 8E).
Description:—Terrestrial, with short to long, at least to 36 cm long, horizontal, usually subterranean, isotomously
branching shoots, from which stiffly erect, aerial shoot systems arise. Aerial shoots to 40 cm tall including erect
subterranean divisions, to 5 times dichotomous. Subterranean divisions with pale appressed leaves, 2–4 mm thick
including leaves. Aerial shoots stiffly erect, homophyllous, terete, equally thick throughout, or slightly tapering, 4–6(–
8) mm thick including leaves. Stems of aerial shoots 1.5–3 mm thick excluding leaves, partly to completely concealed
by leaves, brownish to bright red, with smooth to somewhat rugose epidermis. Leaves borne in alternating, irregular
whorls of 4–5, these 1.5–3 mm apart, forming 8–10 regular to irregular longitudinal ranks, arcuate-appressed to closely
appressed, abaxially convex and rounded or with a prominent, long decurrent veinal ridge, with or without a slightly
prominent basal swelling (air sac), adaxially concave to slightly convex, linear-lanceolate to lanceolate, 4–6(–7) ×
1.2–1.8(–2) mm, upward often reduced to 3.5–4 mm long, red tinged to entirely red, with smooth to unevenly rugose
and sclerified margins, amphistomatic. Sporangia 1–1.3(–1.5) mm wide.
FIGURE 41. A. Phlegmariurus hohenackeri (Hammen & González 1337, COL), growth habit. A1. Close-up of sporangiate divisions.
Distribution:—Southern Colombia to northern Bolivia.
Habitat:—Wet and boggy páramo and jalca, elev. 3000–3400 m.
Notes:—Usually an easily recognizable species of marshes and wet depressions of the lower páramos with a soft
substrate where the subterranean runner-shoots are well developed. When the species grows on more solid substrates,
e.g. open soil in land slides and road cuts, epiterranean creeping shoots with well developed, patent-ascending leaves
replace the runner shoots, and make the distinction from slender forms of Phlegmariurus crassus more subtle. In such
cases the difference of leaf curvature, leaf size, and the number of leaf orthostichies are helpful characters.
Specimens examined:—Cauca: Macizo Colombiano, Páramo de las Papas, around Laguna Cusiyaco, 3360 m,
Barclay & Juajibioy 5980 (GH). Volcán Puracé, Laguna San Rafael, 3300 m, Cleef et al. 590 (COL). Nariño: Near
summit of Cord. Oriental, above Río de Potreros, headwaters of Río Tellez, 3200–3400 m, Ewan 16540 (NO, UC, US).
Cumbal, Volcán Chiles, 28.1.2008, Narvaez 737 (FMB)
Phlegmariurus hystrix (Herter) B.Øllgaard (2012b: 16).—Fig. 42.
Urostachys hystrix Herter (1954: 120).—Lycopodium hystrix (Herter) Morton (1964: 72).—Huperzia hystrix (Herter) Holub (1985: 73).
Type:—ECUADOR: Imbabura, Cordillera Oriental, below camp east of Volcán de Cayambe, on wet, nearly bare, volcanic rock,
10400 ft., Drew E-330 (holotype US).
Description:—Plants ascending to erect, massively robust, mature plants forming large clumps, without basal,
prostrate-ascending, rooting shoots, to 30 cm tall, to 60 cm long, to 5 times dichotomous. Shoots homophyllous, almost
equally thick throughout, (8–)13–23 mm in diameter including leaves, sometimes tapering to 8–13 mm in diameter
upward, sporangiate from 6–15 cm above the base and upward. Stems excluding leaves (3–)5–10 mm thick at the base,
sometimes slightly tapering, almost completely concealed by leaves. Leaves uniform throughout, or slightly smaller
upward, densely crowded, borne in more or less regular, often oblique or obscure, alternating whorls of (5–)8–10, these
1–1.5 mm apart, forming (10–)16–20 often obscure longitudinal ranks, patent-ascending to imbricate, with the apical
third or sometimes the whole leaf strongly upward curved and terminating in a short, slender, recurved, whip-like
tip, broadly triangular-lanceolate to ovate-lanceolate, broadest shortly above the base, (5–)7–10 × 1.5–2.5(–3) mm,
evenly tapering or slightly acuminate, fleshy, with a wide prominent veinal ridge adaxially and abaxially, often almost
quadrangular in the apical half, with or without a short basal swelling abaxially, not or slightly decurrent, usually
strongly red or orange tinged in the apex and margins, with smooth to slightly irregularly rugulate margins. Sporangia
2–2.5 mm wide.
Distribution:—Southern Colombia and Northern Ecuador.
Habitat:—Pioneer on road banks and land slides, or in moss cushions on rocky outcrops in wet páramos, elev.
2890–3200 m.
Notes:—This species is among the largest in the páramo region. When fully deloped it forms large clumps of almost
solid, 2 cm thick shoots. Phlegmariurus hystrix is most closely related to P. ulixis (Colombia), and P. llanganatensis
(Ecuador).
Specimens examined:—Cauca: Nevado del Huila, NE slope, ascending by quebrada Verdua, 3500 m, Rangel
et al. 1454 (PSO, US). Nariño: Páramo of Cerro San Francisco, above Córdoba, 2890–3200 m, Ewan 16244 (COL,
NO, US). Mun. La Cruz, slopes of Volcán Dña. Juana, 3550–3600 m, Mora 3333 (PSO). Mun. Puérres Vereda el
Rosal, 3280 m, Baca et al. Y 329 (PSO). Mun. Tangua, Vereda Las Piedras, Páramo Las Piedras, 3100–3500 m, Baca
et al. Y 083 (PSO). Mun. Santiago, San Antonio de Bellavista, Páramo del Bordoncillo, 3250–3450 m, Ramírez 5165
(PSO). Mun. Pasto, Cord. Centro-oriental, Microcuenca Las Tiendas, inicio del Páramo del Bordoncillo, 3450–3510
m, Ramírez et al. 10573 (PSO). Mun. Guachucal-Cumbal, Páramo del Infernillo, ridge toward Volcán Cumbal, 3700
m, Stancik 2692 (PSO).
Phlegmariurus idroboi B.Øllgaard sp. nov.—Fig. 43.
Diagnosis: Apparently closely related to Phlegmariurus brevifolius, but larger in all parts, with leaves uniformly ca. 4 × 5 mm, consistently
wider than long.
Type:—COLOMBIA: Huila/Cauca: Macizo Colombiano, Páramo de Las Papas, cerros y alrededores de la laguna de La Magdalena, 3530
m, 16. Oct. 1958, Idrobo, Pinto & Bischler 3330 (holotype COL).

FIGURE 42. A. Phlegmariurus hystrix (Ewan 244, COL), growth habit. A1. Close-up of sporangiate division.
Description:—Plants erect or ascending, stiff and robust, forming small clumps, without basal creeping and rooting
shoots, about 25 cm tall, to at least 5 times dichotomous. Shoots homophyllous, almost equally thick throughout, 10–
12 mm in diam. including leaves. Stems excluding leaves 5–6 mm thick, concealed by leaf bases, sporangiate from ca.
15 cm above the base and upward. Leaves almost uniform throughout, densely crowded, borne in regular, alternating
whorls of 4, these 1–1.5 mm apart, forming 8 longitudinal ranks, perpendicular to slightly recurved, with straight to
upward curved apex, broadly ovate to broadly suborbicular-cordate, broadly acute to almost obtuse, 3.5–4 × 4–5 mm,
stiffly coriaceous, red tinged, abaxially concave, with sharply prominent vein, with sharply prominent, short-decurrent
basal swelling (air sac), with flat to slightly revolute, strongly sclerified, darker and somewhat translucent, slightly
rugose margins. Sporangia ca. 2 mm wide, spores normal.
Habitat:—Páramo, at 3530 m.
Notes:—The epithet honours Dr. Jesús M. Idrobo Muñoz of the National University of Colombia for his important
collections of Colombian Lycopodiaceae.
Aditional specimens examined (paratypes): Colombia. Nariño: Volcán Azufral, 18 May 1876, André 3220 (F,
NY).
FIGURE 43. A. Phlegmariurus idroboi (Idrobo, Pinto & Bischler 3330 (COL), growth habit. A1. Close-up of terminal sporangiate
divisions.
Phlegmariurus josesantae B.Øllgaard sp. nov.—Fig. 44.
Type:—COLOMBIA: Antioquia:, Mun. de Urrao, Páramo de Frontino, sitio conocido como “La Virgen”, 3760 m, 13 de Marzo 1986, José
Santa 1192 (holotype AAU).
Diagnosis: Resembling Phlegmariurus dianae, but differs by is larger size in all parts, lanceolate to ovate-lanceolate leaves with leaf
margins minutely rugose-papillose by individually protruding marginal and submarginal cells.
Description:—Plants erect or ascending, usually forming large clumps, to at least 4 times dichotomous, without basal,
creeping and rooting shoots, sometimes more than 20 cm tall. Shoots homophyllous, almost equally thick throughout,
10–15 mm in diameter including leaves. Stems excluding leaves 3–4 mm thick, usually concealed by leaf bases,
except in basal divisions, sporangiate from 8–15 cm above the base and upward, the stem surface sparsely to densely

papillose. Leaves almost uniform throughout, or gradually shorter and wider upward, usually densely crowded in
upper divisions, borne in rather regular alternating whorls of 4–5, these 1.5–2.5 mm apart, forming 8–10 longitudinal
ranks, loosely imbricate or ascending to patent or slightly reflexed, lanceolate (in basal divisions) to ovate-lanceolate,
sometimes with a subcordate base, acute, the apex straight or slightly upward curved, (1.5–)2–3 × 6–8 mm, herbaceous
to subcoriaceous, slightly lustrous, abaxially concave with prominent vein at the lamina base, upward usually convex
with slightly to strongly prominent vein, sometimes convex nearly throughout, with prominent to sharply prominent,
short to long-decurrent basal swelling (air sac), margins flat to slightly involute or sometimes slightly and narrowly
revolute, minutely and densely rugose-papillose by individually protruding marginal and submarginal cells, adaxially
flat to somewhat concave, or with slightly prominent vein, hypostomatic. Sporangia 1.5– 2 mm wide.
FIGURE 44. A. Phlegmariurus josesantae (Santa 1192, AAU), growth habit. A1. Close-up of basal divisions. A2. Close-up of sporangiate
divisions.
Distribution:—Endemic to N Colombia
Habitat:—The type was growing in a thin vegetation cover on basaltic rock, apparently in the company of P.
dianae. Elevation 3400–3760 m.
Notes:—This species is named in honour of José Santa and his important contribution to the documentation and
knowledge of Colombian Lycopodiaceae.
Additional specimens examined (paratypes): Antioquia: Mun. Urrao, Páramo de Frontino, 3580 m, Santa 1191
(AAU), 3580 m, Santa 1189 (AAU), 3450 m, Santa 1178 (AAU), 3500 m, Santa 1181 (AAU). Páramo Frontino, Near
Llano Grande, 3450 m, Boeke & McElroy 274a (NY). Mun. Urrao, Jaiperá, Vereda El Chuscal, Páramo de Frontino,
Llanogrande, 3000–3390 m, Callejas, R. et al. 7631 (AAU).
Phlegmariurus lignosus (Herter) B.Øllgaard (2012b: 16).—Fig. 45.
Lycopodium lignosum Herter (1909a: 46).—Urostachys lignosus (Herter) Nessel (1939: 104).—Huperzia lignosa (Herter) Holub (1985:
74). Type:—COLOMBIA: Pamplona, Las Cruces, 3050 m, Funck & Schlim 1474 (holotype P00559276 [with Herter’s annotation];
isotypes BM, BR; G is a mixed collection including P. serpentiformis).
FIGURE 45. A-A2. Phlegmariurus lignosus (Killip 17967, F) Growth habit.—A3. Phlegmariurus lignosus (Stancik & Medina 2466,
AAU), close-up of sporangiate divisions.
Description:—Plants erect or ascending from a creeping and rooting base, often forming large clumps, often 5–7
times dichotomous, to 25(–35) cm tall. Shoots homophyllous, or with leaves of basal divisions smaller, gradually
becoming larger upward, almost equally thick throughout, 3–5(–8) mm in diameter including leaves. Stems excluding

leaves (2–)3–4 mm thick, almost completely concealed by leaf bases, except in some basal divisions, sporangiate from
5–10 cm above the base and upward. Leaves of creeping base and basal part of aerial shoots small 1–2 mm, gradually
becoming larger upward, densely crowded throughout, borne in regular alternating whorls of 4–5, these 0.6–1 mm
apart, forming 8–10 longitudinal ranks, usually stiffly ascending to patent, or sometimes recurved, usually straight
or with a slightly upward curved apex, broadly triangular-ovate to cordate-orbicular, obtuse to acute and slightly
apiculate, 2–2.5(–3) × 1.5–2(–2.5) mm, coriaceous, usually red tinged, abaxially concave with prominent vein, with
prominent, short decurrent basal swelling (air sac), with flat or slightly revolute, smooth to slightly uneven margins,
adaxially with slightly prominent vein. Sporangia 1–1.5 mm wide.
Distribution:—Endemic to the central and northern Andes of Colombia.
Habitat:—Most collections indicate rocky páramo habitats, elev. 3100–4200 m.
Notes:—This species is unusual in the group og Phlegmariurus brevifolius because the basal divisions are
differentiated into short-creeping and rooting, and aerial, erect and spore-producing shoots. However, the creeping
shoots are not always included in collections. The creeping basal divisions with their small leaves may reflect the
stressful initial growth conditions in rocky habitats before proper rooting is acquired.
Specimens examined:—Boyacá: Mun. Susacón, Mattoral and rocky páramo on the margin of Río Susacón,
20 km passing Susacón—Soata, 3150 m, Stancik & Medina 2466 (AAU). Páramo de Guina, E slope of Quebrada
Los Colorados, 3300 m, Cleef 9758 (AAU, U). Mun. Guicán, Nevado del Cocuy, Corralitos, 3900 m, Bischler 2879
(G). Huila: La Plata, 3000 m, Sneidern 2202 (G). Norte de Santander: Pamplona, páramo de San Urban, 4115 m,
Linden 1235 (BM, BR, K, P), syntype. Norte de Santander/ Cesar: 20 km S of Abrego, Las Jurisdicciones (Cerro de
Oroque), 3700–3960 m, 19–21 May 1969, García-Barriga 19745 (COL, MA, US). Jurisdicciones, Cerro de Oroque,
3700–3900 m, García Barriga & Jaramillo 20655 (U). Santander: Laguna de Cunta, edge of Páramo de Santurbán,
3880 m, Killip & Smith 17967 (F, GH, NY, US). Páramo de Santurbán, near Vetas, 3950–4160 m, Killip & Smith 17476
(COL, GH, MA, S, US). E Cord. Páramo de Las Coloradas, above La Baja, Killip & Smith 18465 (GH, K, US). Mun.
Toná, Serranía de Santurbán, road Berlín-Vetas, páramo de Calaverita, 3690 m, Santa & Escobar 1096 (AAU).
Phlegmariurus llanganatensis (B.Øllg.) B.Øllgaard (2012b: 16).—Fig. 46.
Huperzia llanganatensis B.Øllgaard (1988: 35). Type: ECUADOR. Tungurahua: Páramo de Soguillas, near las Torres de Llanganati,
3850—4000 m, Øllgaard & Holm-Nielsen 38748 (AAU holotype, GH, NY, QCA isotypes).
Illustration: Øllgaard 1988: Fig. 13B.
Description:—Plants ascending to erect, robust, sparsely branched plants without prostrate-ascending, rejuvenating
shoots, to 30 cm tall, or to 80 cm long, to 4 times dichotomous. Shoots homophyllous, 12–17 mm in diam. incl. leaves,
sometimes slightly tapering upward, sporangiate from 10–15 cm above the base and upward. Stems excluding leaves
(3–)5–9 mm thick at the base, often concealed by leaves, with shallow to prominent decurrent ridges from the leaf
bases. Leaves uniform throughout, or slightly shorter and broader upward, somewhat spaced to densely crowded,
borne in more or less regular, often oblique or obscure, alternating low spirals or whorls of (5)7–8, 1–2(–3) mm apart,
forming (10–)14–16 often obscure longitudinal ranks, spreading or slightly recurved and sigmoid to ascending or
loosely imbricate, with the apical part upward curved, usually terminating in a short, slender whip-like tip, ovate-
lanceolate to broadly ovate, broadest in the lower third, (1.5–)2–4 × 4–7 mm, evenly tapering to acuminate, fleshy,
with a broad, somewhat prominent veinal ridge adaxially and abaxially, with a very short, prominent, slightly decurrent
basal swelling abaxially, with red tinged apex and margins, margins smooth to irregularly erose. Sporangia 1.5–2 mm
wide.
Distribution:—Northern Ecuador, Southern Colombia. Wet páramos, elev. 3100 m.
Habitat:—Wet páramos, elev. 3100–4000 m
Notes:—
Closely related to Phlegmariurus hystrix and P. ulixis, but differing by the the shorter, wider leaves. It seems
intermediate between P. hystrix and one of the broad-leaves species in the P. brevifolius group. The sole Colombian
collection is referred to this species with some doubt, being smaller than the average for the species.
Specimens examined:—Nariño: Mun. Pasto: Páramo de Quillinsayaco, 3100 m, Santa & Buitrago 928 (AAU).
FIGURE 46. A. Phlegmariurus llanganatensis (Santa & Buitrago 928, AAU), growth habit. A1. Close-up of terminal sporanguate
division.
Phlegmariurus myrtuosus (Spring) B.Øllgaard (2012b: 17).—Fig. 15B, 47.
Lycopodium myrtuosum Spring (1849: 9).—Urostachys myrtuosus (Spring) Nessel (1927: 387).—Huperzia myrtuosa (Spring) Trevisan
(1874: 248). Type:—COLOMBIA. Magdalena: Santa Marta, Purdie s.n. (holotype K; isotype BM).
Lycopodium lellingeri Rolleri (1978: 143).—Huperzia lellingeri (Rolleri) Holub (1985: 74). Type:—VENEZUELA. Tachira: limestone
outcrops of Páramo de Tamá, near Colombian-Venezuelan border, 3045–3475 m, Steyermark 57362 (holotype US; isotypes MO,
NY).
Description:—Plants erect or ascending, stiff and robust, forming small to large clumps, without basal, prostrate-
ascending, rooting shoots, at least to 25 cm tall, at least to 5 times dichotomous. Shoots homophyllous, almost equally
thick throughout, 9–12 mm in diameter including leaves. Stems excluding leaves 4–6 mm thick at the base, slightly
tapering upward, usually concealed by leaf bases, sporangiate from 5–10 cm above the base and upward. Leaves almost
uniform throughout, densely crowded, or sometimes more spaced in basal divisions, borne in regular, alternating

whorls of 4–5, these 1.5–3 mm apart, forming 8–10 longitudinal ranks, ascending or perpendicular, slightly to strongly
recurved, broadly lanceolate to elliptic or ovate, acute, 4–5 ×2–2.5 mm, stiffly papery to subcoriaceous, green,
abaxially flattened or slightly concave, with slightly prominent to obscure vein, with slightly prominent or indistinct
basal swelling (air sac), with flat to slightly revolute, rugose margins, sometimes red-tinged at apex. Sporangia 1–1.5
mm wide.
Distribution:—Colombia and Venezuela.
Habitat:—Páramos, elev. 2900–3700 m.
Notes:—Exposed individuals approaching the aspect of Phlegmariurus brevifolius, with short, stiff, less fleshy,
perpendicular leaves, were described as Lycopodium lellingeri. Shaded individuals approach the aspect of a compact
P. brongniartii and have longer, softer, recurved-arching leaves.
Specimens examined:—Caldas: Cord. Central, Páramo de Las Letras, ca. 1 mile SW of Letras, elev. 3700 m,
Barclay & Juajibioy 6250 (GH).
FIGURE 47. A. Phlegmariurus myrtuosus (Venezuela, Luteyn et al. 5940, NY), growth habit. A1. Terminal sporangiate divisions.
Phlegmariurus ocananus (Herter) B.Øllgaard (2012b: 17).—Fig. 48.
Lycopodium ocananum Herter (1909a: 45).—Urostachys ocananus (Herter) Nessel (1939: 101).—Huperzia ocanana (Herter) Holub
(1985: 75). Type:—COLOMBIA. Norte de Santander: Ocaña, Schlim 472 (lectotype P-00559163, designated by Øllgaard 1989: 59;
isolectotypes B, BR),.
Phlegmariurus arcturi (Herter) B.Øllgaard (2012b: 12).—Urostachys arcturi Herter (1953: 118). .—Lycopodium arcturi (Herter) C.Morton
(1964: 72). Type: COLOMBIA: Cundinamarca, Haught 5705, (US, holotype, COL, isotype).
Urostachys arthurii Herter (1954: 114).—Lycopodium arthurii (Herter) Morton (1964: 72).—Huperzia arthurii (Herter) Holub (1985:
70). Type:—COLOMBIA. Bolivar: San Cristóbal, Bro. Apollinaris & Bro. Arthur 152 (holotype US 537755).
Urostachys trachyloma Herter (1954: 113).—Huperzia trachyloma (Herter) Rolleri & Deferrari (1988: 157).—Lycopodium trachyloma
(Herter) Morton (1964: 73). Type:—COLOMBIA. Cundinamarca: Choachi, January 1927, Bro. Apollinaris 1 (holotype US
1499319).
FIGURE 48. A. Phlegmariurus ocananus (Santa 1029, AAU) growth habit. A1. Close-up of terminal sporangiate divisions.
Description:—Plants with age forming cespitose tufts, sometimes to 15 cm wide, 5–25 cm tall, usually homoblastic,
sometimes with short creeping (usually not rooting) shoots, at least to 5 times dichotomous. Aerial shoots 5–12(–15)
mm in diameter at the base, including leaves, uniform throughout or narrower, to 5–8(–10) mm in diameter upward.
Aerial stems 1.5–2.5(–4) mm thick (dried) excluding leaves near the base, upward narrower, ca. 1–2(–3) mm thick, stem
surface usually smooth. Leaves of aerial shoots borne in usually regular alternating whorls of 4–5(–6), these usually
1–2 mm apart, forming (6–)8–10(–12) often indistinct longitudinal ranks, homophyllous or gradually reduced upward,
ascending to appressed, usually upward curved (often patent in shaded divisions), leaves of basal divisions 5–7(–10)
× 1–1.2(–2.2) mm, subulate to lanceolate, distally and in sporangiate divisions gradually becoming lanceolate to ovate
(–ovate-cordate, sometimes slightly acuminate), abaxially usually strongly convex or with a prominent vein (dried) or
somewhat concave at the ± perpendicular base, 3–5 × 1–1.7(–2) mm, slightly lustrous to pruinous, adaxially concave,
the apex short-acute to narrowly acuminate and usually upward curved, rarely red tinged (in strongly exposed habitats
at the highest altitudes), leaf margins smooth or usually rugulate by protruding cells, hypostomatic or amphistomatic.
Sporangiate leaves usually with a prominent base, causing a perpendicular appearance of the very leaf base. Sporangia
ca. 1–2 mm wide.
Distribution:—Andes of Venezuela and Colombia.
Habitat:—Páramo, often relatively dry and rocky (2900–)3000–3950 m, at lower altitudes also in shrubby and
grassy páramos.
Notes:—Urostachys moritzii Nessel (1940: 160) is based on rather poor material indicated to be from Venezuela,
may be a synonym of Phlegmariurus ocananus. Urostachys orionis Herter (1949: 53) is a superfluous nomenclatural
synonym for that name, represented by relatively small plants with relatively short and wide, ascending leaves .
An incompletely understood species or species complex, highly variable in size. Some of its features are intermediate
between those of the present , and those of the P. brevifolius group. Some individuals (e.g. Sota 6276 (MER), López-
Figueira 12012 (US), Cuatrecasas et al. 28383 (US) appear to retain juvenile morphology (longer narrower leaves)
in all or in some branches, more mature in other branches.
Specimens examined:—Antioquia: Mun. Urrao, Páramo de Frontino, 3550 m, Santa 1180, 1187 (AAU). Mun.
Urrao, Páramo de Frontino, 3800 m, Santa 1194 (AAU). Mun. Ituango, road El Retiro-Cerro Paramillo, Cuadros 5014,
5019 (MO). Boyacá: Mun. Aquitania, road Sogamoso Pajarito, páramo de Las Cintas, 3200 m, Santa & Escobar 1124
(AAU, HUA ); ibid., 3030 m, Santa & Escobar 1122 (AAU). Mun. Duitama, Páramo de La Rusia, 3500 m, Santa &
Buitrago 883 (AAU). Mun. Duitama, Páramo de La Rusia, La Rusia, 3600 m, Santa & Escobar 1085 (AAU). Páramo
de La Rusia, NW-N of Duitama, Serrania Peña Negra, cerro 1 km ESE of Laguna Agua Clara, 3950 m, Cleef 7436
(AAU). Páramo de Guina, Sta Rosita, E slope of Valle Q Los Colorados, 3300 m, Cleef 9743 (AAU). Mun. Saboya,
Páramo de Saboya, 3000 m, Stancik 1058 (AAU). Mun. Chinavita, Cerro Mamapacha, 2900 m, Stancik 1610 (AAU).
Páramo de Pisva, road Socha—La Punta, km 69, between El Cardón and El Cadillal, 3560 m, Cleef 4646 (COL).
Cauca: Méndez, NE of Silvia, Cord. Central, 3000–3100 m, Haught 5100 (US). Cesar: Páramo de Sabana Rubia,
3250 m, Cuadros 3728 (MO). Cundinamarca: Guadalupe, 3100 m, Apollinaire & Arthur 176 (US). Guadalupe, 3200
m, Haught 5670 (COL). Páramo de Choachi, near Bogotá, 3200–3300 m, Pennell 2242 (NY). Páramo de Sumapáz,
Chisacá, 3660 m, Cleef 4913 (AAU). Mun. Zipaquirá, road to Pacho ascending to Alto del Chamicero, 3000–3200 m,
Santa 1017 (AAU). Mun. Zipaquirá, alto del Chamicero, 3200 m, Santa 1020 (AAU, QCA). Mun. La Calera, páramo
de Palacio, 3500–3800 m, Santa 1029 (AAU). Meta: Páramo de Sumapáz, Cubarral, surroundings of Laguna de la
Guitarra, Franco & Rangel 194 (COL). Santander: Mun. Cerrito, Páramo del Almorzadero, 3500 m, Santa & Escobar
1108 ( AAU, HUA ). Páramo del Almorzadero, 4000 m, Vareschi 4217 (VEN). Valle: Páramo de Pan de Azúcar,
3300–3700 m, Espinal & Ramos 2424 (MEDEL).
Phlegmariurus pachyskelos B.Øllgaard (2016a: 268).—Fig. 49.
Type:—COLOMBIA. Huila: Huila-Cauca, Macizo Colombiano, páramo de Las Papas, Cerros y alrededores de la laguna La Magdalena,
3530 m, 16 October 1958, Idrobo, Pinto & Bischler 3298 (holotype COL; isotype NY).
Description:—Plants erect or ascending, stiff and robust, forming small to large clumps, without basal, prostrate-
ascending, rooting shoots, to more than 40 cm tall, to 5 times dichotomous. Shoots homophyllous, almost equally thick
throughout, 10–15 mm in diameter including leaves, or sometimes tapering to 8 mm in diameter in terminal divisions.
Stems exclcluding leaves 8–10 mm thick at the base (dried), sometimes tapering upward to 5–6 mm, usually concealed
by leaf bases, usually minutely scabrate, sporangiate from 10–25 cm above the base and upward. Leaves almost uniform
throughout, densely crowded, or sometimes more spaced in basal divisions, borne in regular, alternating whorls of 5–6,
these 1.5–2.5 mm apart, forming 10–12 longitudinal ranks, perpendicular to usually sharply recurved-reflexed, with
straight to recurved apex, broadly ovate or triangular-ovate to broadly suborbicular-cordate or triangular-cordate, short-
acute to almost obtuse, sometimes slightly sigmoid, 3.5–5(–6) × (2.5–)3–4(–5) mm, stiffly coriaceous, green, abaxially
concave, with prominent vein, with somewhat prominent, flattened, short-decurrent basal swelling (air sac), adaxially
with prominent vein, slightly revolute, strongly sclerified, darker and somewhat translucent, smooth to minutely erose
margins. Sporangia 1.5–2 mm wide.
Habitat:—High páramos. Elev. 3100–4300 m.
FIGURE 49. A. Phlegmariurus pachyskelos (Idrobo, Pinto & Bischler 3298, COL), growth habit. A1. Close-up of sporangiate division.
Notes:—Appears to be a giant form of Phlegmariurus serpentiformis (the stem 8–10 mm thick,dried), with which
it shares the snake-like, scale-covered aspect.
Specimens examined:—Boyacá: Nevado del Cocuy, alto valle de Las Lagunillas, 4000–4300 m, Cuatrecasas
1508 (F, US). Cauca: Cordillera Central, W slope, Headwaters of Río Palo, high páramos between Quebrada de San
Paulino and Quebrada del López, Lagunilla de Las Casitas, 3700 m, Cuatrecasas 19116 (F, GH, US). Huila/Cauca:
Páramo de Las Papas, near headwaters of Río Magdalena, 3100–3500 m, Camilo s.n. (US). Meta: Páramo de Sumapaz,
Cerro Nevado de Sumapaz, 4000 m, Cleef 1531 (AAU, U)

Phlegmariurus rufescens (Hooker) B.Øllgaard (2012b: 18).—Fig. 50C.
Lycopodium rufescens Hooker (1837: t. 36).—Huperzia rufescens (Hooker) Trevisan (1874: 249).—Urostachys rufescens (Hooker) Nessel
(1939: 103). Type:—ECUADOR: Surucuchu [as »Purucuchu«], Jameson s. n. (holotype K).
Lycopodium mirum Vareschi (1958: 317). Type:—VENEZUELA. Mérida: Sierra Nevada de Mérida, Lago Anteojos, Pico Bolívar, 4150
m, 03 April 1958, Vareschi & 6867 (holotype VEN 42919).
Illustrations:—Øllgaard (1988: Fig. 12D).
Description:—Plants erect or ascending, forming small clumps, to 15(–20) cm tall, usually 2–4(–6) times dichotomous,
without basal, creeping and rooting shoots. Shoots almost homophyllous, almost equally thick throughout, (6–)7–8(–
9) mm in diameter including leaves. Stems excluding leaves 2–3 mm thick, almost completely concealed by leaf
bases, sporangiate from 5–10 cm above the base and upward. Leaves densely crowded throughout, or sometimes more
spaced in basal divisions, borne in rather regular, alternating whorls of 3–4, these ca. 1 mm apart, forming 6–8 usually
distinct longitudinal ranks. Leaves of basal, non-sporangiate divisions ascending to patent, broadly oblong-lanceolate
to ovate, appearing obtuse due ti an upward curved acute apex, gradually wider upward, 2.5–3.5 × 1.5–2 mm, abaxially
rounded or with slightly prominent vein, with evenly, broadly decurrent base. Leaves of upper, sporangiate divisions
loosely imbricate or ascending to perpendicular, rarely reflexed, sometimes slightly sigmoid, broadly ovate to almost
orbicular-cordate, acute to obtuse, (2–)2.5–3.5 × 2–3(–3.5) mm, softly to firmly coriaceous, usually red tinged, with
the widest part abaxially concave to convex, with usually strongly prominent vein at least in the basal half, with a
distinct, short-decurrent basal swelling (air sac), adaxially usually with slightly prominent veinal ridge, usually with
upward curved tip, with usually narrowly sclerified, usually dark and somewhat translucent, smooth to rugose margins.
FIGURE 50. A. Phlegmariurus sellifolius (Ecuador, Holm-Nielsen et al. 5914A, AAU), growth habit. A1. Close-up of sporangiate division.
—B. Phlegmariurus sellifolius, (Ecuador, Asplund 9740, S) growth habit. B1. Close-up of sporangiate division.—C. Phlegmariurus
rufescens (Ecuador, Holm-Nielsen et al. 5914, AAU), growth habit. C1. Close-up of sporangiate division. C2. Two sporophylls. Del. B.
Johnsen.
Distribution:—Andean Venezuela, Colombia to Southern Ecuador Vargas Gaviria, Buitrago & Esquivel (2018)
recorded this species from Páramo de Anaime, Dept. Tolima.
Habitat:—Exposed wet páramos, elev. 3400–4100 m.
Notes:—Closely related to Phlegmariurus brevifolius but smaller, with relatively thin stems, and forming small
individuals with relatively few branches.
Specimens examined:—Boyacá: Páramo de La Rusia, NW-N de Duitama Serrania de Peña Negra, hoya de la
Laguna Agua Clara, 3800 m, Cleef 7065 (AAU, U). Cauca: Cord. Central, E slopes of Páramo del Puracé, around
Laguna de San Rafael, 3320 m, Cuatrecasas & Willard 26300 (COL, US). Magdalena: Sierra Nevada de Santa Marta,
cabeceras de Río Ancho, Páramo de Macotama, Barclay & Juajibioy 6969 (GH, MO).
Phlegmariurus schlimii (Herter) B.Øllgaard (2012b: 18).—Fig. 51.
Lycopodium schlimii Herter (1909a: 45).—Urostachys schlimii (Herter) Nessel (1939: 91).—Huperzia schlimii (Herter) B.Øllgaard (1987:
169). Type:—COLOMBIA. Guajira: Prov. de Río Hacha (Pico Hacha), Sierra Nevada, 11000 p., Schlim 882 (holotype B; isotypes
BONN [Nessel 157], BR, G, K, P).
FIGURE 51. A. Phlegmariurus schlimii (Cuatrecasas & Romero Castañeda 24645 (COL, US), growth habit.—A1. Phlegmariurus
schlimii (Ranghel & Cleef 1001 (AAU). Close-up of sporangiate divisions.

Description:—Plants erect to ascending, to 30 cm tall, to 6 times dichotomous. Shoots gradually heterophyllous,
ca. 20–30 mm in diameter including leaves in basal or juvenile divisions, upward gradually narrowing to 5–10 mm
thick, including leaves. Stem 6–10 mm thick excluding leaves (dried) in basal divisions, striate by decurrent leaf
bases, upward gradually narrowing to 2.5–4 mm thick, glossy red-orange, upward usually completely concealed by
imbricate leaves. Leaves of basal divisions borne in irregular alternating whorls of 4–5, these 2–3 mm apart, forming
8–10 irregular longitudinal rows throughout, upward more closely set, ca. 1–1.5 mm apart. Leaves of basal divisions
and juvenile plants, ascending to recurved, coriaceous, glossy, 10–22 × 2–2.5(–3) mm, flat or with slightly revolute
margins, with distinct vein, abaxially usually with to rugose bands of slightly depressed stomata (dried), Leaves of
upper, sporangiate divisions, closely imbricate, glossy, coriaceous, reddish tinged, lanceolate to triangular-lanceolate
strongly convex abaxially, apically somewhat involute, clasping the sporangia, 5–11 × 1.5–2 mm, with smooth margins.
Sporangia ca. 2 mm wide.
Distribution:—Colombia: Sierra Santa Marta; Vargas Gaviria, Buitrago & Esquivel (2018) recorded this species
from Páramo de Anaime, Dept. Tolima.
Habitat:—Páramo, bushy prairies, and thickets, 3300–3900 m.
Notes:—A highly local and striking endemic. The species is placed in P. crassus group with doubt. It resembles
the Brazilian P. ruber and may represent a parallel adaptation derived from elements in the P. brongniartii group.
Specimens examined:—Magdalena: Sierra Nevada de Santa Marta, near headwaters of Río Ancho, Páramo
de Macotama, 3840 m, Barclay & Juajibioy 6926, 6928 (GH). Near origin of Río Yebosimeina, 3900 m, White &
Alverson 576 (HUA, NY). Drainage of Río Donachuí, rocky slopes of Chitebo, 3450–3550 m, Cuatrecasas & Romero
Castañeda 24645 (COL, US). Sierra Nevada de Santa Marta, SE slopes, Drainage of Río Donachuí, Melloaca, 3320–
3260 m, Cuatrecasas & Castañeda 24447 (COL, US). Transecto del Buritaca, Filo La Cumbre, 3800 m, Ranghel &
Cleef 1001 (AAU, U).
Phlegmariurus schmidtchenii (Hieronymus) B.Øllgaard (2012b: 19).—Fig. 52
Lycopodium schmidtchenii Hieronymus (1905: 570).—Urostachys schmidtchenii (Hieron.) Nessel (1939: 103).—Huperzia schmidtchenii
(Hieron.) Holub (1985: 76). Type:—COLOMBIA: Monte Ruiz ad Rio Guali, 1882, Schmidtchen s. n. (holotype B n.v., holotype
fragment in NY).
Lycopodium goudotii Herter (1909a: 47).—Urostachys goudotii (Herter) Nessel (1939: 106).—Huperzia goudotii (Herter) Holub (1985:
73).—Phlegmariurus goudotii (Herter) B.Øllgaard (2012b: 15). Type:—COLOMBIA: Páramo de Usaquén, Jan. 1844, Goudot 3
(holotype P; isotype fragment BONN-Nessel 202).
Description:—Plants erect or ascending, usually forming small to large clumps, at least to 5 times dichotomous,
without basal, and rooting shoots, to 30 cm tall. Shoots homophyllous, almost equally thick throughout, 5–8(–12) mm
in diameter including leaves. Stems excluding leaves 3–5 mm thick, almost concealed by leaf bases except in basal
divisions, sporangiate from 6–10 cm above the base and upward. Leaves almost uniform throughout, usually densely
crowded throughout, borne in rather regular alternating whorls of 5–6 (–8), these 1–1.5 mm apart, forming 10–12(–16)
longitudinal ranks, (patent or) often strongly recurved, often sigmoid, ovate-lanceolate to oblong-lanceolate, acute
to slightly acuminate, 2–4(–5) × 1.2–2 mm, coriaceous, green or red tinged, hypostomatic, abaxially concave with
prominent vein in basal part of lamina, with prominent to sharply prominent, short to long-decurrent basal swelling
(air sac), with flat to slightly revolute, narrowly sclerified, smooth to slightly rugulate margins, adaxially with slightly
prominent vein. Sporangia 1–1.5 mm wide.
Distribution:—Endemic.
Ramírez Padilla & Macias Pinto (2007) reported this species from Dept. Cauca.
Habitat:—Páramo, elev. 3000–4000 m
Notes:—Resembles Phlegmariurus serpentiformis and P. sellifolius, but differs in the consistently oblong-
lanceolate, acute to acuminate leaves.
Specimens examined:—Caldas: Nevado del Ruíz, Pachón 10 (COL). Nevado del Ruiz, road Nevado–Manizales,
3750 m, Santa & Hoyos 688 (AAU). Mun. Villamaría, road Manizales–Nevado del Ruiz, 3740 m, Santa, J. et al. 957
(AAU) . Mun. Villamaría, road Manizales–Bogotá, 5 km above road to Nevado del Ruiz, 3480–3500 m, Arbeláez,
A. et al. 455 (MO). Cundinamarca: Páramo de Chisacá, 3800–3960m, Bernal 238 (COL). Paramo de Guasca, rich
bank near cloud forest, 3200 m, Gillett & Jaramillo 16587 (COL, MO). Cord. Oriental, mun. Fomeque, Paramo de
Chingaza, cerros vecinos a La Laguna, 3500 m, Huertas & Camargo 6589A (COL). Cercanías del Páramo de Cruz
Verde, 3000 m, Romero Castañeda 2533 (COL). La Calera, Páramo de Palacio, carretera Adelante de La Mina, 3700
m, Uribe 6702 (COL). Tolima: Road Mariquita–Manizales, Alto de Letras, 3300 m, Santa, J. & L. A. de Escobar 855
(AAU, OCA)
FIGURE 52. A. Phlegmariurus schmidtchenii (Huertas & Camargo 6589 p.p., COL), growth habit.—B. Close-up of sporangiate division
(Schmidtchen s. n., holotype fragment NY).

Phlegmariurus sellifolius (B.Øllg.) B.Øllgaard (2012b).—Fig. 50A-B.
Huperzia sellifolia B.Øllgaard (1988: 68). Type:—ECUADOR: Prov. Carchi, Road Tulcán–Maldonado, at the pass, km 38, 4000 m,
Øllgaard & Balslev 8432 (holotype, AAU).
Urostachys compactus (Hook.) Nessel f. minor Herter (1954: 119–120). Type: COLOMBIA: Nariño: Páramo of Cerro San Francisco,
above Córdoba, Cord. Oriental, 3100 m, Ewan 16243 (holotype US, isotype NO).
Illustrations:—Øllgaard (1988: Fig. 12C)
Description:—Plants erect or ascending, usually forming large clumps, often 5–7 or more times dichotomous, without
basal, creeping and rooting shoots, often more than 25 cm tall. Shoots homophyllous, almost equally thick throughout,
5–8 mm in diameter including leaves. Stems excluding leaves 2.5–3(–4) mm thick, almost completely concealed by
leaf bases, except in basal divisions, sporangiate from 6–20 cm above the base and upward. Leaves almost uniform
throughout, usually densely crowded throughout, borne in rather regular alternating whorls of 4–5, these 0.6–1.5 mm
apart, forming 8–10 longitudinal ranks, loosely imbricate or ascending to patent or sharply reflexed and appressed
to the stem, often distinctly sigmoid, broadly lanceolate to ovate, ovate-triangular or elliptic, acute or obtuse, (1.5–
)2–2.6(–3.5) × (1.2–)1.5–2(–2.4) mm, coriaceous, often brightly red tinged, abaxially concave with prominent vein
in basal part of lamina, to convex at the apex or sometimes convex nearly throughout, with prominent to sharply
prominent, short to long-decurrent basal swelling (air sac), with flat to slightly involute, narrowly sclerified, smooth to
slightly uneven margins, adaxially with slightly to distinctly prominent vein. Sporangia 1–1.5 mm wide.
Distribution:—Andes of Southern Colombia to Northern Peru.
Habitat:—Exposed moist páramos, elev. 2800–4100 m.
Notes:—Phlegmariurus sellifolius is is often found in company of the closely related P. rufescens. It is smaller
than the latter species in all parts, has narrower leaves, and usually has has more leaf ranks. It usually forms large
clumps with numerous branches, while P. rufescens usually is rather sparsely branched, and forms small clumps
only.
Specimens examined:—Cauca: Macizo Colombiano, Páramo de Las Papas, El Boquerón and surroundings of
Laguna La Magdalena, 3500 m, Idrobo & Barclay 4046 (COL, G, US). Cordillera Central, Puracé, 3450 m, Sneidern
1695 (S). Páramo de Moras, between Mozoco and Pitayó, Tierras Adentro, 3000–3500 m, Pittier 1414 (US). Huila/
Cauca: Macizo Colombiano, Páramo de Las Papas, surroundings of Laguna La Magdalena and Santiago, 3530–3630
m, Idrobo et al. 3332, 3345 (COL). Idrobo, Pinto & Bischler 3283 (COL). Nariño: Páramo de Quilinsayaco between
Pasto and Sibundoy, Barclay 4559 (COL, MO). Mun. Pasto, páramo of Quillinsayaco, 3100 m, Santa & Buitrago
926, 927 (AAU, HUA). Mun. Guachucal–Cumbal. Páramo Infernillo toward Volcán Cumbal, grass páramo, 3700 m,
Stancik 2691 (AAU). Alta cuenca del río Putumayo, filo de la Cordillera entre El Encano y Sibundoy, páramo de San
Antonio de Bordoncillo, 3250 m, Cuatrecasas 11767 (COL). Mun. Pasto, Correg. La Laguna, surroundings of Alto
Zapallurco, 3250 m, Ramírez & Salas 10816 (PSO).
Phlegmariurus serpentiformis (Herter) B.Øllgaard (2016b: 269).—Fig. 53.
Lycopodium serpentiforme Herter (1909a: 46).—Urostachys serpentiformis (Herter) Nessel (1939: 104).—Huperzia serpentiformis
(Herter) Rolleri & Deferrari (1988: 156). Type.—COLOMBIA. Norte de Santander: Ocaña, Schlim 341 (holotype P00559275,
annotated as type by Herter; isotypes BR, K).
Description:—Plants erect to ascending, stiff and robust, forming small to large clumps, to more than 40 cm tall, to
6 times dichotomous. Shoots homophyllous, almost equally thick throughout, 6–8(–10) mm in diameter including
leaves, or sometimes tapering to 5 mm in diameter in terminal divisions. Stems exclcluding leaves (3–)5–7 mm thick
at the base (dried), sometimes tapering upward to 3–5 mm, usually concealed by leaf bases, often minutely scabrate,
sporangiate from ca.10 cm above the base and upward. Leaves almost uniform throughout, densely crowded, or
sometimes more spaced in basal divisions, borne in regular, alternating whorls of (4–)5(–6), these 1–1.5 mm apart,
forming (8–)10(–12) longitudinal ranks, (perpendicular to) usually sharply recurved-reflexed, with straight to recurved
or slightly sigmoid apex, broadly ovate or triangular-ovate to broadly suborbicular-cordate or triangular-cordate, the
outer (reflexed) face often wider than long, almost obtuse to short-acute or apiculate, sometimes slightly sigmoid,
2–4 × 1.5–2.5 mm, stiffly coriaceous, green, abaxially concave, with prominent vein, with a very distinct prominent,
basal swelling (air sac), adaxially with prominent vein, margins slightly revolute, sclerified, smooth to minutely erose.
FIGURE 53. A. Phlegmariurus serpentiformis (Rangel 1657A, COL), growth habit. A1. Close-up of sporangiate divisions.
Distribution:—Endemic to central Colombia.

Habitat:—Páramo, elev. 3200–4200 m.
Notes:—Phlegmariurus serpentiformis appears like a half-size P. pachyskelos, with the the leaves usually closely
reflexed and appressed like the scales of a snake. It is closely related also to Phlegmariurus schmidtchenii and to P.
sellifolius and P. brevifolius. The latter three are often hard to separate from P. serpentiformis.
There are three specimens of Schlim 341 in P, one from Herbier Christ (not annotated by Herter) is mixed,
containing four specimens, the one to the right of the middle is identical to the holotype, the three others are P. lignosus.
Another specimen from Herbier Houllet (not annotated by Herter) is P. lignosus.
Specimens examined:—Cundinamarca: Páramo de Palacio (La Calera), 3600 m, Hammen 388, 390 (COL).
Páramo de Palacio, Hda. La Siberia, 3200–3400 m, Mora 942 (COL). Páramo de Palacio, Lagunas de Buitrago y
alrededores, 3600 m, Cleef & t’Hart 2328 (AAU, U). Páramo de Palacio, Lagunas de Siecha, 3700 m, Cleef & Jaramillo
4042 (U). Mun. Guasca, W slope, 3300 m, Sarmiento 180 (COL, MO). Páramo de Siberia, 3500 m, Schneider 1192
(COL). Páramo de Palacio, 10 km E of La Calera, 20 km NE of Bogotá, Tryon & Tryon 6041 (COL, UC). Región del
Sumapáz, 2 km from división a la carretera a San Juan, 3960 m, Rangel 1657A (COL). Paramo de Sumapaz, Cuchilla
La Ramona entre Andabobos y cabeceras del Río San Juan, 4000 m, Cleef 1626 (U). Paramo de Chisacá, alrededores
de lagunas de Chisacá, hoya del río Magdalena, 3700–3800 m, Pinto & Hernández 514 (COL). Laguna de Chisacá y
alrededores, 3900–4200 m, Bischler 1510 (COL, G). E cordillera S of Usme, Paramo de Chisacá, 4020 m, Barclay &
Juajibioy 6212 (MO). Páramo de Palacio, Quebrada de Casarreales, páramo, 3450 m, Cuatrecasas et al. 25630 (COL).
Páramo de Chisacá, Laguna Negra, 3660–3720 m, Cuatrecasas & Jaramillo 25907 (COL). Valle del río Tunjuelo en
Páramo de Chisacá, alrededores de Laguna Grande, 3400–3600 m, Murillo et al. 771 (COL). Páramo de Chisacá,
alrededores de la Laguna, Murillo & Fayad 44 (COL). Cruz Verde, S of Bogotá, Holton s.n. (K, NY). Mun. Fomeque,
Páramo de Chingaza, cerros vecinos a La Laguna, 3500 m, 3500 m, Huertas & Camargo 6589B (COL). Meta: W
side of Nevado del Sumapáz, 1300 m, Little 7013 (COL). Cord. Oriental, mun. Fomeque, Páramo de Chingaza, cerros
vecinos a La Laguna, 3500 m, Huertas & Camargo 6589 p.p. (COL). Páramo de Sumapaz, Cerro Nevado del Sumapaz,
3950 m, Cleef 1337 (AAU, US). Páramo de Sumapaz, Cerro Nevado del Sumapaz, superpáramo, 4015 m, Cleef 7735
(AAU). Páramo de Sumapaz, Cerro Nevado del Sumapaz, 3575 m, Cleef 8228 (U).
Phlegmariurus sphagnicola B.Øllgaard (2016b: 270).—Fig. 54.
Type:—COLOMBIA: Magdalena: Sierra de Perijá, E of Manaure: Sabana Rubia, 3000–3100 m, 6–8 November 1959, Cuatrecasas &
Romero-Castañeda 25036 (holotype COL)
FIGURE 54. A. Phlegmariurus sphagnicola (Cuatrecasas & Romero Castañeda 25036, COL), growth habit. A1. Close-up of young
individual with slender basal part.
Description:—Plants erect, or with a short creeping and rooting prostrate shoot, forming loose clumps of sparsely
branched (paludicolous?) individuals, to twice ramified, to at least 35 cm tall. Shoots homophyllous and equally thick
throughout or slightly tapering upward, 8–15 mm in diameter including leaves near the base, sporangiate from 10–15
cm above the base and upward. Stems excluding leaves 1.2–2 mm thick at the base, sometimes slightly tapering
upward nearly completely concealed by the leaves. Leaves borne in irregular, alternating whorls of 6 to 8, these 1–1.5
mm apart, forming 12–16 indistinct longitudinal ranks, densely crowded, ascending or usually appressed, spreading
in shaded shoots and in parts covered by sphagnum, almost straight to slightly upward curved at leaf apex, linear-
lanceolate to linear, 7–10 × 0.8–2 mm in basal divisions, upward often reduced to 4–6 × 0.5–1.5 mm, adaxially concave
or flat, with slightly prominent vein, abaxially rounded (or with a median ridge, dried) with slightly prominent, long
decurrent basal swelling (air cavity), amphistomatic, with smooth, narrowly sclerified margins, lustrous and slightly to
strongly red tinged. Sporangia 1.0–1.3 mm wide.
Distribution:—Sierra Nevada de Santa Marta, 3000–3570 m.
Habitat:—Known from Sphagnum bogs and wet slopes in páramo, 3000–3600 m.
Specimens examined:—Magdalena: Sierra Nevada de Santa Marta, alrededores de cabeceras de Río Sevilla,
3570 m, Barclay & Juajibioy 6639 (GH, MO). Upper Mamancanaca Valley, S side of the major peaks peat bed, 3650
m, Cabot 17 (GH).
Phlegmariurus transilla (Baker) B.Øllgaard (2012b: 19).—Fig. 55.
Lycopodium transilla Baker (1877: 168).—Urostachys transilla (Baker) Nessel (1939: 106).—Huperzia transilla (Baker) Holub (1985:
77). Type:—ECUADOR: Volcán de Cayambe, al nivel de las nieves perpetuas 4600 m, Sodiro s. n. (holotype K; isotypes BP, NY
fragment, UC).
Illustrations:—Øllgaard (1988: Fig. 11A).
Description:—Plants erect or erect from a decumbent base, very robust, to 50 cm tall, to 1 m long, to 4 times
dichotomous. Shoots homophyllous, 20–30 mm in diameter including leaves. Stems excluding leaves 12–15 mm
thick, slightly tapering upward, red tinged to bright red, not concealed by the leaves, terete or with short decurrent
ridges from the leaf bases, sporangiate nearly from the base. Leaves uniform throughout, borne in alternating whorls
of 6, patent to ascending, ovate–ovate-lanceolate, distinctly narrowed at the very base, 8–11 × 3.5–4.5 mm, adaxially
convex to concave, dark glaucous green, rigid and brittle, with abaxially obscure to slightly prominent vein, with flat
to revolute, smooth to minutely and irregularly crenulate margins; sporangia ca. 1.5 mm wide.
Distribution:—Colombia and northern Ecuador.
Habitat:—Scrub páramo and wet páramo, usually with some shelter, elev. 3800–4100 m.
Notes:—Baker’s original spelling of the epithet is maintained, even if Sodiro (1883, 1893) corrected it to
‘trencilla’. Baker followed Sodiro’s spelling on the type sheet in K, and on an isotype in BP.
This is one of the largest and most impressive clubmosses in the Andean páramos, unlikely to be mistaken for
any other species. The Colombian specimens deviate from the Ecuadorian ones by their narrower and slightly shorter
leaves.
Sodiro (1883: 112) reported the use of this species as an effective medicine improving blood circulation in the
liver, and causing menstruation.
Specimens examined:—Chocó: Mun. Nóvita, W slope of range Tatamá–Pico Belisario (in honor of President
B. Betancour), S ridge, 4100 m, Torres et al. 2444 (AAU, COL). Macizo de Tamaná, NW of Cerro Ventanas, 3860 m,
Torres 1854 (COL). Santander: Mun. Coromoro, Laguna de Fiquera o Jequera, Páramo de Carnicerías, sources of
Quebrada Coromoro, 3850 m, Jaramillo & van der Hammen 4370 (COL).
Phlegmariurus ulixis (Herter) B.Øllgaard (2012b: 20).—Fig. 56.
Urostachys ulixis Herter (1954: 115).—Lycopodium ulixis (Herter) C.Morton (1964: 73).—Huperzia ulixis (Herter) Holub (1985: 77).
Type:—COLOMBIA. Cauca: Cordillera Central, W slope, headwaters of Río Palo, high páramos between Quebrada de San Paulino
and Quebrada del López, Lagunilla de Las Casitas, 3700 m, Cuatrecasas 19085 (holotype US; isotypes F, GH).
Description:—Plants ascending to erect, robust, mature plants forming large clumps, homoblastic, to 30 cm tall, to 6
times dichotomous. Shoots homophyllous, almost equally thick throughout, 12–18 mm in diameter including leaves,
sometimes tapering to 8–10 mm in diameter upward, sporangiate from 10–15 cm above the base and upward. Stems
excluding leaves 4–6 mm thick at the base, sometimes slightly tapering, almost completely concealed by leaves.
Leaves uniform throughout, or slightly smaller upward, densely crowded, borne fairly regular, alternating whorls of
5–6 (–7), these 1–2 mm apart, forming 10–12(–14) often obscure longitudinal or somewhat sloping ranks, patent-
ascending to imbricate, with the apical third or sometimes the whole leaf strongly upward curved, acute or terminating
in a short, slender sometimes recurved whip-like tip, broadly triangular-lanceolate to ovate-lanceolate, widest shortly
above the base, 5–8 × 2–3 mm, evenly tapering or slightly acuminate, somewhat fleshy, with a prominent veinal ridge
abaxially, flattened thoughout or the apex slightly quadrangular, with a small basal swelling abaxially, not or slightly
decurrent, green to strongly red or orange tinged at the apex and margins, with slightly to coarsely irregularly rugulate
margins, hypostomatic. Sporangia ca. 1.5–2 mm wide.
Habitat:—Páramo, 2475–3800 m.
FIGURE 55. A. Phlegmariurus transilla (Mejia & van der Hammen 4370, COL) growth habit. A1. Close-up of terminal sporangiate
division.
Notes:—This species resembles Phlegmariurus hystrix (Ecuador and Southern Colombia). The type specimens of
P. ulixis are young individuals with leaves flatter and wider than in P. hystrix. Generally, the leaves of P. ulixis are flat
thoughout and not apically subquadrangular as usual in P. hystrix.
Specimens examined:—Cauca: Carretera Belalcázar–Tacueyó, km 35, 2475 m, Rangel 2582 (COL). Páramo
de Las Papas, surroundings of Laguna Cusiyaco, 3500 m, Barclay & Juajibioy 6026 (GH). Huila/Cauca: Macizo
Colombiano, páramo de Las Papas y cerros alrededores de la Laguna de La Magdalena, 3530 m, Idrobo et al. 3117 (COL
p.p.), 3318 (COL, VEN). Nariño: Páramo de Bordoncillo, 3250 m, Espinal 1011 (COL, US). Páramo de Quillinsayaco,
between Pasto and Sibundoy, Barclay 4551 (MO). Páramo de Quillinsayaco, 3100 m., Santa & Buitrago 928 (AAU,
COL, HUA, MEDEL ). Putumayo, Páramo de Quilinsayaco, 3200 m, Hagemann & Leist 1342 (PSO). Valle: Cord.
Central, W slope, headwaters of Ríos Tuluá and Bugalagrande: Páramo de Las Vegas, 3600–3800 m, Cuatrecasas
20337 (S, US p.p.).
FIGURE 56. A. Phlegmariurus ulixis (Cuatrecasas 19085, F), growth habit.—A1-A2: Phlegmariurus ulixis (Barclay 6026, GH), close-
up of middle- and terminal sporangiate divisions.

Excluded names
Huperzia apolinari-mariae (Nessel) Holub (1985: 70). The type of this name is indicated to be Colombian, but belongs
to Huperzia selago (L.) Martius & Schrank, which is unknown in tropical America. So another case of nesselian
mislabelling is the likely explanation.
Huperzia aristei (Nessel) Rolleri & Deferrari (1988: 155). No material matching the type of Urostachys aristei
Nessel is known from Colombia. Considering its similarity to Bolivian and Argentinean collections of Phlegmariurus
sotae (Rolleri) B.Øllgaard, and being familiar with Nessel’s frequent lack of precision, I assume that the type label is
erroneous.
Phlegmariurus saururus (Lam.) B.Øllgaard (2012b: 283), in the Neotropics this species is limited to the Andes of
Peru and further south.
Urostachys Kupperi Nessel (1934: 180). The Colombian type resembles a form of Phlegmariurus crassus, but I
cannot decide about its identity.
Acknowledgements
The working facilities and generous assistance, particularly by Finn Borchsenius and Birgitte Bergmann, for this
project in Herbarium AAU of the institute of Bioscience, and the Science Museums of Aarhus University are gratefully
acknowledged. I am grateful to Weston Testo for inspiring and helpful correspondence, and to Henrik Balslev for
encouraging conversations.
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Phytotaxa 426 (1): 001–105 ISSN 1179-3155 (print edition)

Synopsis of the genus Phlegmariurus (Lycopodiaceae) in

Accepted by Marcus Lehnert: 24 Oct. 2019; published: 16 Dec. 2019 1

FIRST PUBLISHED IN 2019 BY

© 2019 Magnolia Press

ISSN 1179-3155 (print edition)

2 • Phytotaxa 426 (1) © 2019 Magnolia Press ØLLGAARD

Keywords: Lycopodiaceae, Phlegmariurus, Colombia

Group of Phlegmariurus myrsinites

6 • Phytotaxa 426 (1) © 2019 Magnolia Press ØLLGAARD

8 • Phytotaxa 426 (1) © 2019 Magnolia Press ØLLGAARD

10 • Phytotaxa 426 (1) © 2019 Magnolia Press ØLLGAARD

Phlegmariurus ericifolius (Presl) B.Øllg.

Phlegmariurus myrsinites (Lamarck) B.Øllgaard (2012b: 17).—Fig. 6.

12 • Phytotaxa 426 (1) © 2019 Magnolia Press ØLLGAARD

Notes:—Phlegmariurus myrsinites is related to P. phylicifolius. Forms of the latter, described as Lycopodium

Phlegmariurus phylicifolius (Poiret) B.Øllgaard (2012b: 17).—Fig. 1A.

Phlegmariurus subulatus (Poiret) B.Øllgaard (2012b: 19).—Fig. 1B.

14 • Phytotaxa 426 (1) © 2019 Magnolia Press ØLLGAARD

Group of Phlegmariurus acerosus

Phlegmariurus acerosus (Swartz) B.Øllgaard (2012a: 480).—Fig. 7A.

16 • Phytotaxa 426 (1) © 2019 Magnolia Press ØLLGAARD

Phlegmariurus curvifolius (Kunze) B.Øllgaard (2012b: 14).—Fig. 8C.

Description:—Delicate to extremely delicate plants, flaccidly pendulous, to 60 cm long. Shoots gradually

Phlegmariurus filiformis (Swartz) W.H. Wagner (1995: 242).—Fig. 7B.

Phlegmariurus tenuis (Willdenow) B.Øllgaard (2012b: 19).—Fig. 8A–B.

18 • Phytotaxa 426 (1) © 2019 Magnolia Press ØLLGAARD

Phlegmariurus arcuatus (B.Øllg.) B.Øllgaard (2012b: 12).—Fig. 9A.

20 • Phytotaxa 426 (1) © 2019 Magnolia Press ØLLGAARD

26. Phlegmariurus dichotomus (Jaquin) W.H.Wagner (1993: 305).—Fig. 5B.

Phlegmariurus hippurideus (Christ) B.Øllgaard (2012: 480).—Fig. 10A-B.

24 • Phytotaxa 426 (1) © 2019 Magnolia Press ØLLGAARD

Phlegmariurus pithyoides (Schlechtendal & Chamisso) B.Øllgaard (2012b: 17).—Fig. 12B.

Phlegmariurus polycarpos (Kunze) B.Øllgaard (2012b: 18).—Fig. 5A.

26 • Phytotaxa 426 (1) © 2019 Magnolia Press ØLLGAARD

Phlegmariurus wilsonii (Underwood & F.Lloyd) B.Øllgaard (2012a: 481).—Fig. 13.

Description:—Plants erect, arcuate-spreading to pendulous, to 20(–30) cm long. Shoots homophyllous, 1.5–2.5(–3)

Group of Phlegmariurus hartwegianus

Phlegmariurus campianus (B.Øllg.) B.Øllgaard (2012b: 13). Fig. 14A.

28 • Phytotaxa 426 (1) © 2019 Magnolia Press ØLLGAARD

Phlegmariurus echinatus (Spring) B.Øllgaard (2012b: 14).—Fig. 14B.

30 • Phytotaxa 426 (1) © 2019 Magnolia Press ØLLGAARD

Description:—Robust, recurved or pendulous plants, at least to 90 cm long. Shoots homophyllous to gradually

Group of Phlegmariurus lindenii

Phlegmariurus lindenii (Spring) B.Øllgaard (2012b: 16).—Fig. 1C.

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Group of Phlegmariurus taxifolius

Phlegmariurus taxifolius (Swartz) Löve & Löve (1977: 324).—Fig. 16.

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Group of Phlegmariurus linifolius

Phlegmariurus homocarpus (Herter) B.Øllgaard (2012b: 15).—Fig. 17A.

Phlegmariurus killipii (Herter) B.Øllgaard (2012a: 480).—Fig. 12A.

36 • Phytotaxa 426 (1) © 2019 Magnolia Press ØLLGAARD

Phlegmariurus linifolius (Linnaeus) B.Øllgaard (2012a: 480).—Figs. 17–19.

A. Phlegmariurus linifolius var. linifolius—Fig. 17B.

Illustrations: Øllgaard (1988: Fig. 16B).

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Phlegmariurus sarmentosus (Spring) B.Øllgaard (2012b: 18).—Fig. 18B.

40 • Phytotaxa 426 (1) © 2019 Magnolia Press ØLLGAARD

Group of Phlegmariurus brongniartii

Phlegmariurus brongniartii (Spring) B.Øllgaard (2012b: 12).—Fig. 9B.

Phlegmariurus rosenstockianus (Herter) B.Øllgaard (2012b: 18).—Fig. 15C-D.

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