A. I. Jocouand N. F. Brignone - Juncus articulatus in the Southern Cone

FirSt record oF JuncuS articulatuS SubSP.

articulatuS (Juncaceae) For the Southern cone

Flora

Primer regiStro de JuncuS articulatuS SubSP. articulatuS

(Juncaceae) Para la Flora del cono Sur

1

2

Adriel I. Jocou * and Nicolás F. Brignone

Summary

Background and aims: Juncus is the largest and most diverse genus of Juncaceae,

distributed mainly in the temperate zones of both hemispheres. The Southern Cone

ﬂora contains ca. 38 Juncus species (44 taxa including 12 infraspeciﬁc categories).

Sixteen of those species belong to the section Ozophyllum. As a part of our ecologic

studies in wetlands of Patagonia (Argentina) we collected specimens of Juncus that

did not match any of the species currently known to the Southern Cone. The aim

of this contribution is to report for the ﬁrst time the presence of Juncus articulatus

subsp. articulatus for the Southern Cone ﬂora.

M&M: Classical methods in taxonomy were employed. Living and herbarium material,

original descriptions, and type material of Juncus articulatus subsp. articulatus

were studied. Collected materials were deposited in the herbarium ARC of the

Universidad Nacional del Comahue.

Results: Juncus articulatus subsp. articulatus is here described. A distribution map,

photos, and a key to the Southern Cone species of Juncus belonging to the section

Ozophyllum are provided. Also, some ecological and distributional features are

discussed.

Conclusions: The naturalization of Juncus articulatus subsp. articulatus is reported

here for the ﬁrst time in the Southern Cone. From now on, Juncus contributes 39

species to the Southern Cone ﬂora, and 17 to Juncus section Ozophyllum.

1

. Departamento de Biología

Aplicada, Facultad de Ciencias

Agrarias. Universidad Nacional del

Comahue. RN N° 151 km 12,5,

CC 85, CP 8303, Cinco Saltos, Río

Negro, Argentina.

2

. Instituto de Botánica Darwinion

(

IBODA, CONICET-ANCEFN),

Labardén 200, CC 22, CP 1642, San

Isidro, Buenos Aires, Argentina.

*adrieljocou@gmail.com

Citar este artículo

JOCOU, A. I. & N. F. BRIꢀNONE.

020. First record of Juncus

articulatus subsp. articulatus

Juncaceae) for the Southern Cone

flora. Bol. Soc. Argent. Bot. 55:

31-640.

2

(

6

Key wordS

DOI: https://doi.

Flora of Argentina, Juncus, new record, Ozophyllum, South America, wetlands.

org/10.31055/1851.2372.v55.

n4.29938

reSumen

Introducción y objetivos: Juncus es el género más grande y diverso de Juncaceae,

distribuido en zonas templadas de ambos hemisferios. La ﬂora del Cono Sur

contiene ca. 38 especies de Juncus (44 taxones que incluyen 12 categorías

infraespecíﬁcas). Dieciséis de esas especies pertenecen a la sección Ozophyllum.

Como parte de nuestros estudios ecológicos en humedales de la Patagonia

(Argentina), hemos coleccionado especímenes de Juncus que no coincidían con

ninguna de las especies actualmente conocidas en el Cono Sur. El objetivo de esta

contribución es reportar por primera vez la presencia de Juncus articulatus subsp.

articulatus para la ﬂora del Cono Sur.

M&M: Se emplearon los métodos clásicos de taxonomía. Se estudió material vivo y

de herbario, descripciones originales y material tipo de Juncus articulatus subsp.

articulatus. El material coleccionado fue depositado en el herbario ARC de la

Universidad Nacional del Comahue.

Resultados: Se describe a Juncus articulatus subsp. articulatus. Se proporciona

un mapa de distribución, fotografías y una clave de las especies de Juncus

pertenecientes a la sección Ozophyllum presentes en el Cono Sur. Se discuten

algunas características ecológicas y de distribución de la especie.

Conclusiones: Se reporta por primera vez la naturalización de Juncus articulatus

subsp. articulatus en el Cono Sur. Desde ahora, Juncus aporta 39 especies a la

ﬂora del Cono Sur y 17 a la sección Ozophyllum.

Recibido: 15 Ago. 2020

Aceptado: 17 Sep. 2020

PalabraS clave

Flora Argentina, humedales, Juncus, nuevo registro, Ozophyllum, Sudamérica.

Publicado en línea: 13 Nov. 2020

Publicado impreso: 20 Dic. 2020

Editora: Carolina I. Calviño

ISSN versión impresa 0373-580X

ISSN versión on-line 1851-2372

631

Bol. Soc. Argent. Bot. 55 (4) 2020

introduction

As a part of our ecological and ﬂoristic studies

in the wetlands of Patagonia, Argentina (Fernández

Juncaceae Juss. is a cosmopolitan family, et al., 2018; Jocou & Gandullo, 2018; Jocou et al.,

comprising seven genera and ca. 440 species 2018; Gandullo et al., 2019; Jocou & Gandullo,

(Stevens, 2001 onwards; Kirschner, 2002a). Rushes 2020; Jocou et al., 2020), we collected specimens

are mainly distributed in temperate and polar of Juncus that did not match the features to any

regions in both hemispheres, commonly in the of the species currently known to the Southern

subtropics, inhabiting the mountains (Kirschner, Cone. The aim of this contribution is to report for

2

3

002a; Drábková et al., 2006). Juncus L. (ca. the ﬁrst time the presence of Juncus articulatus

15 spp.), and Luzula DC. (ca. 115 spp.) are the subsp. articulatus for the Southern Cone ﬂora, by

most widespread genera in the family, whereas the study of living and herbarium material, original

Marsippospermum Desv. (four species) and descriptions, type material, and complementing

Rostkovia Desv. (two species) are found in New with a distribution map, photos, and a key to the

Zealand, South America and on some southern sect. Ozophyllum for the Southern Cone. Also,

hemisphere islands. Oxychloë Phil. (ﬁve species), some ecological and distributional features are

Patosia Buchenau (one species), and Distichia discussed.

Nees & Meyen (three species) are restricted to the

Andes of South America.

The genus Juncus is distributed mainly in both the materialS and methodS

northern and southern hemispheres, but rare in the

tropics (Balslev, 1996; Kirschner, 2002b; Romero

Previous works on the genus Juncus in the

Zarco, 2010). Two subgenera and 10 sections are region were consulted (Barros, 1945; Barros,

currently recognized (Kirschner, 2002b): subg. 1953; Barros, 1969; Balslev, 1996; Novara, 2009;

Juncus (six sections), and subg. Agathryon Raf. Balslev, 2018; Zuloaga et al., 2019). Classical

(four sections; Table 1). The Southern Cone ﬂora methods in taxonomy were employed. The

(

Argentina, south of Brazil, Chile, Paraguay, taxonomic identity of the collected specimens was

and Uruguay) harbours ca. 38 species, and 12 established following Balslev (1996), Kirschner

infraspeciﬁc taxa of Juncus, summing for a total (2002b), and Romero Zarco (2010). The collected

of 44 taxa (Zuloaga et al., 2019). Sixteen of specimens were also compared with the high

those 38 species belong to the sect. Ozophyllum resolution images of the lectotype kept at L

Dumort. (Table 1). The mainly terete leaves that are herbarium (herbarium acronyms following Thiers,

unitubulose or pluritubulose, and perfectly septate, 2020), and with available descriptions of J.

constitute the synapomorphies of this section articulatus, and the subsp. articulatus (Nilsson

(Kirschner, 2002b).

& Snogerup, 1972; Balslev, 1996; Brooks &

Juncus articulatus L. belongs to sect. Ozophyllum Clemants, 2000; Guofang & Clemants, 2000;

sensu Kirschner, 2002b), and it is considered Kirschner, 2002b; Romero Zarco, 2010). Some

(

an extremely variable species, so its description additional material from P, SI, US and WTU

turning difﬁcult in terms of infraspeciﬁc taxa. In belonging to North American and European J.

this sense, Kirschner (2002b) recognizes only two articulatus subsp. articulatus were also studied

subspecies: subsp. articulatus, and subsp. limosus for comparison. All the specimens collected of

(Vorosch.) Vorosch. The typical subspecies is one J. articulatus subsp. articulatus were deposited

of the most widely distributed taxa in Juncus; it is in ARC. The number indicated for the lectotype

frequent in Asia, Europe, North of Africa, North belongs to the barcode of the specimen.

America, also Australia, New Zealand, South

The morphological study of the specimens was

Africa, and Peru (Balslev, 1996; Kirschner, 2002b; performed under a stereomicroscope Leica EZ4

Romero Zarco, 2010). Juncus articulatus subsp. HD, and the photographs were taken using LAS

limosus is restricted to the Far East, being most EZ 3.4.0. The distribution map for the Southern

common in the Amur and Khabarovsk regions, Cone (Fig. 1) was created in QGIS 3.14 “Pi”

and the adjacent regions of Chinese Manchuria (QGIS Development Team, 2020) using the GPS

(Kirschner, 2002b).

coordinates from the collected specimens.

632

A. I. Jocouand N. F. Brignone - Juncus articulatus in the Southern Cone

Table 1. Summary of subgenera and sections within Juncus, as well as the number of species in each

section and their distributions worldwide, and the number and species per section in the Southern Cone

ﬂora. The * indicates the total number of species within section Ozophyllum, including the new record of

J. articulatus subsp. articulatus.

Number

of species

in the

Number and species in the

Southern Cone Flora

Subgenus

Section

Distribution

world

Forskalina

Kuntze

Distributed in the whole Mediterra-

nean region.

1

not represented

Juncus balticus Willd.; J.

8

:

World-wide distribution. Mainly conglomeratus L.; J. effusus L.;

in Australia and Pacific North J. ﬁliformis L.; J. kleinii Barros; J.

Juncotypus

Dumort.

67

America.

procerus E.Mey.; J. ramboi Barros; J.

uruguensis Griseb.

Agathryon

Raf.

Widespread, with species native 7: Juncus capillaceus Lam.; J.

to all temperate regions except for cordobensis Barros; J. dichotomus

South Africa. Mainly in W and E Elliott; J. homalocaulis F.Muell. ex

North America, Central Asia and Benth.; J. imbricatus Laharpe; J.

Steirochloa

Griseb.

35

temperate South America.

tenuis Willd.; J. venturianus Castillón

Tenageia

Dumort.

W of Mediterranean region; some

species very widespread.

11

1: Juncus bufonius L.

Caespitosi

Cout.

1

6

2

Southern Africa, W North America. 1: Juncus capitatus Weigel

Graminifolii

Engelm.

W North America, southern Africa, 3: Juncus cyperoides Laharpe; J.

2

Australasian region.

marginatus Rostk.; J. planifolius R.Br.

Iridifolii

Snogerup &

Kirschner

10

9

W North America, Eastern Asia.

not represented

Not geographically circumscribed.

It includes species from Europe,

America, Africa and East Asia.

2

: Juncus acutus L.; J. kraussii

Juncus

Hochst.

1

L .

7 * :

J u n c u s

s u b s p .

a r t i c u l a t u s

a r t i c u l a t u s ;

Juncus acuminatus Michx.;

J. andersonii Buchenau; J.

austrobrasiliensis Balslev; J.

brasiliensis Breistr.; J. bulbosus L.;

Juncus

E North America, South America, J. burkartii Barros; J. debilis A.Gray;

SW Europe, the Far East, North J. densiflorus Kunth; J. diemii

Ozophyllum

Dumort.

86

Africa, S Africa, Oceania.

Barros; J. ernesti-barrosii Barros; J.

llanquihuensis Barros; J. micranthus

Schrad. ex E.Mey.; J. microcephalus

Kunth (incl. J. involucratus Steud.

ex Buchenau); J. pallescens Lam.;

J. scheuchzerioides Gaudich.; J.

stipulatus Nees & Meyen

In high mountains of North

Hemisphere and subarctic

regions, centred in the Sino-

Himalaya.

Stygiopsis

[

Grand. ex]

Kuntze

59

not represented

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Bol. Soc. Argent. Bot. 55 (4) 2020

reSultS and diScuSSion

Juncus articulatus L., Sp. Pl. 1: 327. 1753.

subsp. articulatus. TYPE: Herb. A. van Royen,

L Nº 904.145–409 (Lectotype L 0221767 [photo!]

designated by Ferrer-Gallego, 2019). Figs. 2-4.

Herbs, perennial, usually caespitose, 10‒55 cm

tall. Rhizome short, short-creeping, rarely with

nodes elongated, 2–3 mm diam., not swollen. Stems

usually erect, sometimes decumbent, submerged

or growing in moving sand and rooting at nodes,

terete, 1–3.5 mm diam., smooth. Cataphylls usually

1

, rarely 2, maroon to straw-coloured, apex acute

to obtuse. Cauline leaves 2–6, usually 3; leaf

sheath auricles well developed, free part 0.5–2

mm, membranous, apex rounded, scarious; leaf

blade green to straw-coloured, terete, 3–17 cm

long, 0.5–2 mm wide, subacute, unitubulose,

with perfect septa that are conspicuous externally.

Fig. 1. Juncus articulatus subsp. articulatus.

Inflorescences decompound, 2–12 long, 1.5–6 Geographic location of collected specimens in

cm wide, terminal panicles of 6–55, obpyramidal Argentina.

Fig. 2. Juncus articulatus subsp. articulatus. A: Plant on urban irrigation canal in Allen, Río Negro province

(Jocou et al. 2314). B: Inﬂorescence (Jocou et al. 2314). C: Plant on rural irrigation canal in General

Fernández Oro, Río Negro province (Jocou 2317). D: Inﬂorescence (Jocou 2317).

634

A. I. Jocouand N. F. Brignone - Juncus articulatus in the Southern Cone

to hemispheric, 4–10-ﬂowered heads, each head

–4 mm long, 3–9 mm wide; branches spreading,

3

erectopatent; primary bract erect, shorter than the

inﬂorescence, 1.5–4 cm long. Tepals maroon or

straw-coloured, completely brown or with a green

central band, ovate to lanceolate, usually equal,

2

.5–3 mm long, 0.7–1.1 mm wide; outer tepals

sometimes slightly longer, carinate, more or less

narrowly bordered, varying in colour from green

dorsally with darker lateral bands and apex dark

brown or castaneous-coloured, acute or rarely

subobtuse and mucronate; inner tepals obtuse to

acute, mucronate, with distinct scarious margins.

Stamens 6, anthers ca. 1 mm long, equalling

ﬁlaments, rarely slightly longer or shorter. Style

0

.3–0.7 mm long, stigmas to 1.5 mm long. Capsule

exserted 0.5–1.5 mm beyond perianth, trigonous,

ellipsoid or ovoid, imperfectly 3-locular, 3.8–4.2

mm long, castaneous to dark brown, glossy, apex

acute proximal to beak, valves separating and apex

slightly reﬂexed at dehiscence; rostrum ca. 0.3–0.5

mm long. Seeds obovoid, 0.5–0.7 mm long,

reticulate, not tailed, yellowish, one of the ends

dark brown coloured.

Fig. 3. Juncus articulatus subsp. articulatus,

vegetative features. A: Specimens rooting at basal

and upper nodes (Jocou 2316). B: Presence of

cataphylls in the lower area of the stem (Jocou

References. See Kirschner (2002b) for list of

synonyms.

2317). Scales= A: 3 cm, B: 2 cm.

Iconography. Bauhin (1620: 12; 1658: 75, 76);

Brooks & Clemants (2000: 253); Thomé (1886: pl.

1

07); Guofang & Clemants (2000: Fig. 42 [9-12]);

Kirschner (2002b: 253, Fig. 109); Morison (1699: 1988; Brooks & Clemants, 2000; Kirschner, 2002b;

sect. 8, t. 9, Fig. 1); Nilsson & Snogerup (1972: Ferrer-Gallego, 2019); 0–3700 m (Guofang &

[

136]-137, Fig. 74).

Clemants, 2000; Brooks & Clemants, 2000).

InArgentina, Juncus articulatus was found growing

Distribution. Widespread: North of Africa, Asia, on river banks (Neuquén River), also in ﬂooded or

Europe, North America and South America, also in wet soils of man-made irrigation canals (Fig. 2A-

Australia, New Zealand, and South Africa (Nilsson D). It has not been observed in drainage channels.

&

Snogerup, 1972; Brooks & Clemants, 2000; Along irrigation canals, this subspecies grows with

Kirschner, 2002b; Romero Zarco, 2010). Bidens pilosa L. var. pilosa (Asteraceae), Epilobium

In South America, Juncus articulatus were ﬁrst ciliatum Raf. (Onagraceae), Juncus microcephalus

reported by Balslev (1996) only for Peru. The Kunth (Juncaceae), Lilaeopsis occidentalis J.M.

presence in the Southern Cone is reported here for Coult. & Rose (Apiaceae), Plantago australis Lam.

the ﬁrst time, growing in Argentina, particularly in subsp. australis (Plantaginaceae), Polypogon viridis

Neuquén and Río Negro provinces (Fig. 1).

(Gouan) Breistr (Poaceae), and Triglochin striata

Ruiz & Pav. (Juncaginaceae). On river banks, it

Habitat. Juncus articulatus subsp. articulatus grows with Plantago australis subsp. australis,

inhabits various moist and wet natural or man-made Cyperus eragrostis Lam. var. eragrostis (Cyperaceae),

places, margins of watercourses, usually on acid Eleocharis bonariensis Nees (Cyperaceae),

soils, often behaving like calcicole (Grime et al., E. radicans (Poir.) Kunth (Cyperaceae), Mentha

635

Bol. Soc. Argent. Bot. 55 (4) 2020

Fig. 4. Juncus articulatus subsp. articulatus, reproductive features. A-B: Flowered-head (Jocou et al. 2314,

Jocou 2317, respectively). C: Colour variability of the tepals, and different maturity stages of capsules (Jocou

&

Minué 2313, Jocou et al. 2314, Jocou 2316, Jocou 2317, respectively). D: Details of capsule and remnants

of the ﬂower (Jocou & Minué 2313). E: Seeds (Jocou 2316). Scales= A-D: 1 mm, E: 0.25 mm.

pulegium L. (Lamiaceae), Polypogon australis articulatus is an extremely variable species with

Brongn. (Poaceae), and P. elongatus Kunth var. a high morphological diversity (Ferrer-Gallego,

elongatus (Poaceae). We observed that this taxon 2019), so its infraspeciﬁc delimitation is difﬁcult,

roots even at the last node of the stem, generating and further studies are needed (Kirschner, 2002b).

new plants and promoting colonization of the habitat Following Kirschner (2002b), the characters of the

through asexual reproduction (Fig. 3A).

specimens collected and examined for this paper

belong to the subsp. articulatus, with tepals to 3

Observations. The water level dynamics and mm long, equal, completely brown or with a green

soil nutrients promote a high morphological central band (never totally green, Fig. 4 A-D), and

plasticity (Grime et al., 1988). As expected, Juncus anthers equalling the ﬁlaments.

636

A. I. Jocouand N. F. Brignone - Juncus articulatus in the Southern Cone

Vogt et al. (2004) conﬁrmed that the dispersal of naturalized in Argentina, according to Pyšek et al.

Juncus seeds occurs through water, and then they (2004).

are deposited in the sediments. Consequently, the

presence of this taxon on the banks of the Neuquén

River, and along the irrigation canals in the Río concluSionS

Negro province may not be a mere coincidence.

The irrigation water comes from artiﬁcial reservoirs

The naturalization of Juncus articulatus

of the Neuquén River, being able to transport the subsp. articulatus is reported for the ﬁrst time in

seeds (Fig. 4E) over long distances, promoting the Southern Cone. From this contribution, the

the distribution of the species along other rivers circumscription of Juncus sect. Ozophyllum to the

and irrigation canals, among other interconnected Southern Cone is here expanded, including Juncus

aquatic environments. In this case, the populations articulatus. Summing up, Juncus comprises 39

found were perfectly established at a distance of species to the Southern Cone ﬂora, and 17 in sect.

more than 30 km from each other. Consequently, Ozophyllum.

and adding the capacity of its seeds to remain viable

for at least 12 years (Geoffrey et al., 1996; Smith articulatus subsp. articulatus around the world,

Brock, 1997; Roberts & Marston, 2000), Juncus further studies might reveal additional occurrences

Due to the widespread distribution of Juncus

&

articulatus subsp. articulatus can be considered as in the Southern Cone.

Key to the Southern Cone species of Juncus belonging to the section Ozophyllum

1

. Leaves bitubulose to pluritubulose.

J. bulbosus

'. Leaves unitubulose.

2

. Stamens usually 3.

3

. Plants rhizomatous; rhizomes creeping, with internodes usually at least 0.5 cm long or longer, or

rhizomes ascending-branching.

4

. Flower heads dark brown.

5

. Capsule more or less equalling the perianth.

J. brasiliensis

5

'. Capsule clearly longer than the perianth.

J. austrobrasiliensis

4

'. Flower heads stramineous, greenish or light castaneous, sometimes with reddish tinge.

6

. Outer tepals conspicuously longer than the inner ones.

J. densiﬂorus

'. Outer tepals more or less equalling the inner ones, sometimes slightly longer or shorter.

J. micranthus

6

3

'. Plants densely caespitose with a very short rhizome, or rhizome short and densely branched or

shortly horizontal, with internodes abbreviated.

7

. Capsule conspicuously longer than the perianth.

J. debilis

7

'. Capsule shorter than, as long as, or slightly longer than the perianth.

8

. Tepals light brown to greenish; capsule subeaqual to slightly exceeding the perianth.

J. acuminatus

8

'. Tepals castaneous to brown; capsule shorter than the perianth.

637

Bol. Soc. Argent. Bot. 55 (4) 2020

9

. Capsule obtuse.

J. ernesti-barrosii

J. burkartii

'. Capsule more or less acute.

2

'. Stamens usually 6.

1

0. Inﬂorescence usually 1 headed, rarely 2 – 3 headed, 2 – 5 (– 8)-ﬂowered each.

1. Lower bract usually exceeding the inﬂorescence; tepals longer than 3.5 mm; style 0.7 – 1.5

mm long.

1

J. scheuchzerioides

11'. Lower bract usually slightly shorter than the inﬂorescence; tepals shorter than 3.5 mm; style

to 0.7 mm long.

J. stipulatus

1

0'. Inﬂorescence 2 or more headed, or if 1-headed then more than 11-ﬂowered.

1

2. Plants densely caespitose; rhizome very short or short and densely branched.

1

3. Capsule shorter than the perianth.

1

4. Flower head peduncles 0.45 mm thick or thicker; tepals rigid.

J. pallescens

1

4'. Flower head peduncles 0.4 mm thick or thinner; tepals soft.

J. microcephalus

1

3'. Capsule exceeding to much exceeding the perianth.

1

5. Capsule light castaneous, rostrum 0.6 – 1.0 mm long; inﬂorescence 8–16-headed, heads

globose, 5–20-ﬂowered, 8-11 mm in diam., ﬂower pedicels to 1.5 mm long; inner tepals

obtuse.

J. andersonii

1

5'. Capsule castaneous to dark brown, rostrum to 0.5 mm long; inﬂorescence 6–55-headed,

heads obpyramidal to hemispheric, 4–10-ﬂowered, 3–9 mm in diam., ﬂower pedicels to

ca. 0.5 mm long; inner tepals acute to subobtuse.

J. articulatus

1

2'. Plants loosely caespitose; rhizome long, creeping.

1

6. Style ca. 1 mm long.

J. diemii

1

6'. Style 0.1 – 0.5 mm long.

J. llanquihuensis

Specimens examined. ARGENTINA. Prov.

Additional specimens examined. CANADA.

Neuquén: Confluencia, Centenario, Balneario Prov. British Columbia: Vancouver Island, near

municipal, 29-XII-2016, Jocou & Minué 2313 (ARC). Lake Shawnigan, 18-VIII-1897, Canby et al.

Prov. Río Negro: General Roca, Allen, 12-XII-2018, 304 (US). Prov. Prince Edward Island: Prince,

Jocou et al. 2314 (ARC); Cinco Saltos, 3-II-2020, Wellington, 26-VII-1952, Erskine 1386 (P). Prov.

Jocou 2318 (ARC); Cipolletti, 12-XII-2018, Jocou et Ontario: Thunder Bay, Mouth of Pine River on

al. 2315 (ARC); General Fernández Oro, 9-I-2020, Lake Superior, Crooks Township, 22-VIII-1952,

Jocou 2316 (ARC); General Fernández Oro, 9-I- Garton 2205 (P). FINLAND. Áland: Lemland,

2020, Jocou 2317 (ARC).

Järsö, recreation area, NE corner of Söderﬁjärden,

638

A. I. Jocouand N. F. Brignone - Juncus articulatus in the Southern Cone

6-VIII-1991, Lampinen & Lampinen 13455 (SI). bibliograPhy

0

Northern Ostrobothnia: Pyhäjärvi, Keliä, in ripa

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STATES. (s. d.), damp bare grounds, 1853-

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1-167. http://www.jstor.org/stable/4393863

1

856, Wright s. n. (US). St. Washington: Clark, BALSLEV, H. 2018. Two new species of Juncus

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(Juncaceae) from South America. Phytotaxa 375:

Rd., 7-XII-2016, Maze CN254 (WTU); Franklin,

97-102. https://doi.org/10.11646/phytotaxa.376.2.3

Hanford National Wildlife Refuge, Wahluke Unit BARROS, M. 1945. Juncáceas argentinas. Holmbergia

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