Part 6. JUNCACEAE 1: Rostkovia to Luzula - Species Plantarum ...

AUSTRALIAN BIOLOGICAL RESOURCES STUDY, CANBERRA 

SPECIES PLANTARUM 

FLORA OF THE WORLD 

Part 6. JUNCACEAE 1: Rostkovia to Luzula 

compiled by Jan Kirschner

© Commonwealth of Australia 2002 

This work is copyright. Apart from any use as permitted under the Copyright Act 1968, no part may be 

reproduced or distributed by any process or stored in any retrieval system or data base without prior 

written permission from the copyright holder. Requests and inquiries concerning reproduction and rights 

should be addressed to the Director, Australian Biological Resources Study, GPO Box 787, Canberra, 

ACT 2601, Australia. 

This work may be cited as: 

J. Kirschner et al., Juncaceae 1: Rostkovia to 

Luzula, Species Plantarum: Flora of the World 

Part 6: 1–237 (2002). 

National Library of Australia 

Cataloguing in Publication entry 

Kirschner, Jan. 

Juncaceae. 1, Rostkovia to Luzula. 

Bibliography. 

Includes index. 

ISBN 0 642 56822 7. 

EXECUTIVE EDITOR 

Anthony E. Orchard 

ASSISTANT EDITORS 

Jennifer Bleyerveen 

Annette J.G. Wilson 

Brigitte Kuchlmayr 

This book is available from: 

Australian Biological Resources Study 

GPO Box 787 

Canberra, ACT 2601 

Australia 

1. Juncaceae. I. Australian Biological Resources Study. 

II. Title. (Series : Species plantarum: flora of the world ; pt. 6). 

584.82

CONTENTS 

Contributors iv 

Species Plantarum Committee v 

Introduction vi 

Preface vii 

Juncaceae 1 

Rostkovia 2 

Marsippospermum 5 

Oxychloë 9 

Patosia 13 

Distichia 15 

Luzula 18 

subg. Marlenia 20 

subg. Luzula 21 

sect. Anthelaea 21 

sect. Atlanticae 37 

sect. Nodulosae 38 

sect. Diprophyllatae 39 

sect. Alpinae 57 

sect. Thyrsanochlamydeae 83 

sect. Luzula 91 

subg. Pterodes 164 

Appendix 188 

Maps 189 

Index 229 

iii

iv 

CONTRIBUTORS TO THIS PART 

Text 

Coordinator & compiler: 

Jan Kirschner, Institute of Botany, Academy of Sciences, CZ-25243 Průhonice 1, Czech 

Republic. 

Advisors to the whole family treatment: 

Sven Snogerup, Botanical Museum, Ö. Vallgatan 18, S-22361 Lund, Sweden. 

Vladimir S. Novikov, Botanical Garden, Moscow State University, Vorobjevy Gory, Moscow 

119899, Russia. 

Teuvo Ahti, Department of Ecology & Systematics, University of Helsinki, P.O. Box 47, 

FIN-00014 Helsinki, Finland. 

Authors (* = Coauthor of the given taxon): 

Henrik Balslev, Herbarium, Building 137, Aarhus University, DK-8000 Aarhus, Denmark 

(Rostkovia*, Patosia, Distichia*). 

Adolf Češka, P.O. Box 8546, Victoria, B.C. V8W 3S2, Canada (Luzula* of Pacific North 

America). 

Janice Coffey Swab, Meredith College, Raleigh, NC 27607-5298, U.S.A. (Luzula* of North 

America). 

Elizabeth Edgar, 32 Harrowdale Drive, Christchurch 4, New Zealand (Luzula* of Australia, 

New Zealand). 

Karmele Garcia-Herran, Université de Neuchâtel, Rue Emile-Argand 11, 2007 Neuchâtel, 

Switzerland (Luzula spicata s.lat.*, Luzula sylvatica s.lat.*). 

Leena Hämet-Ahti, Kotitorpantie 2-E, FIN-00690 Helsinki, Finland (Luzula sect. 

Diprophyllatae*, Luzula sect. Thyrsanochlamydeae*). 

Zdeněk Kaplan, Institute of Botany, Academy of Sciences, CZ-25243 Průhonice 1, Czech 

Republic (Luzula subg. Pterodes). 

Jan Kirschner, Institute of Botany, Academy of Sciences, CZ-25243 Průhonice 1, Czech 

Republic (Rostkovia*, Marsippospermum, Oxychloë* Distichia*, Luzula, generic and 

family treatments). 

Lazaro J. Novara, Herbario MCNS, Facultad de Ciencias Naturales, Universidad National de 

Salta, Buenos Aires 177, 4400 Salta, Argentina (Oxychloë*, Luzula sect. Alpinae*). 

Vladimir S. Novikov, Botanical Garden, Moscow State University, Vorobjevy Gory, Moscow 

119899, Russia (Luzula* of Russia, Central Asia). 

Aaron Wilton, Landcare Research New Zealand Ltd., Canterbury Agriculture & Science 

Centre, P.O. Box 69, Gerald Street, Lincoln, New Zealand (Luzula* of New Zealand). 

Illustrations 

Illustrations prepared for this volume (Plates 2, 13, 17, 19, 20, 36, 37, 38) were drawn by 

Eva Smrčinová, U průhonu 11, CZ-17000 Praha 7, Czech Republic. 

Other illustrations in this work have been reproduced, with permission, from a number of 

sources, which are acknowledged with each plate.

Species Plantarum Steering Committee (as at May 2002) 

G. Achoundong, Herbier National de Cameroon, Yaoundé, Cameroon 

A. Anton, Museo Botánico, Universidad Nacional de Córdoba, Argentina 

W. Berendsohn, Botanischer Garten und Botanisches Museum, Berlin, Germany 

D.E. Boufford, Harvard University Herbaria, Cambridge, Mass., USA 

L. Boulos, Dokki, Cairo, Egypt 

R.K. Brummitt, Herbarium, Royal Botanic Gardens, Kew, UK 

S. Castroviejo, Real Jardín Botánico, CSIC, Madrid, Spain 

A.C. Chikuni, National Herbarium and Botanic Gardens of Malawi, Zomba, Malawi 

M.D. Correa, Universidad de Panamà, Panama 

T.F. Daniel, California Academy of Sciences, San Francisco, Calif., USA 

D.Darnaedi, Kebun Raya Indonesia, Bogor, Indonesia 

P. Dávila, Escuela Nacional de Estudios Profesionales, UNAM, Tlalnepantla, Mexico 

T.S. Filgueiras, Reserva Ecológica do IBGE, Brasilia, Brazil 

E. Forero, Universidad Nacional de Colombia, Bogotá, Colombia 

I. Friis, Botanical Museum and Library, Copenhagen, Denmark 

V. Funk, Smithsonian Institution, Washington DC, USA 

D. Geltman, Komarov Botanical Institute, St Petersberg, Russia 

A.S. George, Kardinya, WA, Australia 

W. Greuter, Botanischer Garten und Botanisches Museum, Berlin, Germany 

Hong D.Y., Chinese Academy of Sciences, Beijing, China 

K. Iwatsuki, Yokohama, Japan 

B. Jonsell, Bergius Foundation, Stockholm, Sweden 

B. Khayota, National Museums of Kenya, Nairobi, Kenya 

J. Kirschner, Academy of Sciences, Pruhonice, Czech Republic 

H.P. Linder, Insitut für Systematische Botanik, Zurich, Switzerland 

P.-A. Loizeau, Conservatoire et Jardin Botaniques, Genève, Switzerland 

J.L. Luteyn, New York Botanic Gardens, Bronx, New York, USA 

K. Marhold, Slovak Academy of Sciences, Bratislava, Slovakia 

M. Moraes, Universidad Mayor de San Andres, La Paz, Bolivia 

P. Morat, Muséum National d'Histoire Naturelle, Paris, France 

N.R. Morin, The Arboretum at Flagstaff, Arizona, USA 

Nguyen T.H., Institute of Ecology & Biological Resources, Hanoi, Vietnam 

H. Ohashi, Tohoku University, Sendai, Japan 

A.E. Orchard, Australian Biological Resources Study, Canberra, ACT, Australia 

R.J. Pankhurst, Royal Botanic Garden, Edinburgh, UK 

M. Sanjappa, Botanical Survey of India, Calcutta, India 

T. Santisuk, Royal Forest Department, Bangkok, Thailand 

Sebsebe Demissew, Addis Ababa University, Addis Ababa, Ethiopia 

J.H. Seyani, Commonwealth Secretariat, London, UK 

G.F. Smith, National Botanical Institute, Pretoria, South Africa 

W.D. Stevens, Missouri Botanical Garden, Saint Louis, Missouri, USA 

W.L. Wagner, Smithsonian Institution, Washington DC, USA 

K. Wilson, Royal Botanic Gardens, Sydney, NSW, Australia 

v

vi 

INTRODUCTION 

Species Plantarum aims to provide in concise format, and with standardised data fields, basic 

taxonomic information on the vascular plants of the world, including accepted names and 

synonyms with bibliographic data, types of names, keys and descriptions from family to 

varietal levels, geographical distributions, ecological information and other related matters, 

and to publish it in both hard copy and electronic form. 

The format of the Species Plantarum is based on that of Flora of Australia, with some 

departures made necessary by the different scale of the project. Initially at least, the series is 

being edited and published for the Species Plantarum Project and IOPI by the Australian 

Biological Resources Study (ABRS), producers of Flora of Australia. 

Treatments are contributed on a voluntary basis. Each part of Species Plantarum is intended 

to provide a complete account of a family, subfamily, large genus or other related taxonomic 

group. While treatments of small families may be shorter, it is intended that contributions 

will, in general, cover at least 50 to 100 species. The taxonomy adopted is that of the author, 

although the family delimitations recommended are initially those of R.K.Brummitt, 

Vascular Plant Families and Genera (1992). The order of taxa within families, genera and 

species in the Species Plantarum is intended to reflect natural relationships, so far as this is 

possible in a linear sequence. 

Maps are provided for each species, or in those cases where infraspecific taxa are recognised, 

for each of the terminal taxa. Distribution maps are based on those in R.K.Brummitt, World 

Geographical Scheme for Recording Plant Distributions ed. 2 (2001), and the 'countries' 

adopted are those of Level 3 and 4 of that work. Description of distribution follows the same 

work, with a two-digit code for regions and a three-letter code for the 'country'. Upper case 

letters for the 'country' indicate native distribution; lower case letters indicate that the taxon 

is only present in that 'country' as an introduced and naturalised plant. If a taxon is extinct in 

a 'country', this is indicated by a dagger. Distribution of species as cultivated plants is not 

included. 

Misapplied and invalid names are, in general, omitted. Journal titles are abbreviated 

according to G.D.R.Bridson & E.R.Smith, Botanico-Periodicum-Huntianum / Supplementum 

(1991). Book titles are abbreviated according to F.A.Stafleu & R.S.Cowan, Taxonomic 

Literature (2nd edn) Vols 1–7, and Supplements (1976–), except that upper case initial letters 

are used for proper names and significant words. Authors of plant names are abbreviated 

according to R.K.Brummitt & C.E.Powell, Authors of Plant Names (1992). 

A separate part, Introduction to the Series, provides a history of the project, a glossary, guide 

for contributors and key to the conventions used in describing distribution. These resources 

will also be available on the World Wide Web, initially through the ABRS site (currently 

http://www.anbg.gov.au/abrs/flora/spplant/spplant.htm) with links from the IOPI site 

(currently http://life.csu.edu.au/iopi/iopihome) and others. 

A.E.Orchard 

Canberra 

May 2002

PREFACE 

This volume is the first of a three-part treatment of the Juncaceae. Parts 2 and 3 are written 

as this goes to press and are scheduled for publication very shortly. 

The co-operative monographic study was completed in the period 1997 to 2000, under grant 

no. 206/97/0936 of the Czech National Grant Agency. Almost all the current experts in the 

taxonomy of the family kindly participated in the project, and their expertise covered both 

the centres of taxonomic diversity and the most intricate groups in the family. The 

monograph thus presents the cumulative knowledge of the Juncaceae at this time. Some 

groups received special attention and their accounts represent new revisions (e.g., Luzula 

sect. Atlanticae, sect. Alpinae, subg. Pterodes) or recent revisions were incorporated (e.g., 

Juncus sect. Caespitosi, in part, and Juncus sect. Juncus). The project benefited from an 

important coincidence: the Juncaceae have been included in recent volumes of important 

regional or continental floras, either published in 1999 or 2000, or prepared for publication; 

the authors of several regional accounts participated in our project (including those from 

Flora of North America, Flora of China, Flora of Australia and Flora Neotropica). Despite 

this, some groups or sections still require a thorough revision in future, and a number of 

questions remain unanswered, particularly in Luzula sect. Luzula in China and Japan, Juncus 

sect. Stygiopsis and Juncus sect. Juncotypus. 

While the text represents the combined work of many authors, the final version was prepared 

by JK, and he accepts responsibility for any errors or missing data. 

In addition to the authors and advisers, many other botanists participated in solving 

numerous problems with old literature, nomenclatural problems and geographical 

distribution. I would like to express my thanks for their generous help. Invaluable technical 

assistance was provided by Ms Věra Matějovičová. Dr R.K.Brummitt of Kew, the Secretary 

of the Flora of the World project, showed unfailing interest, and without his support the 

present volumes would never have come into being. 

Jan Kirschner 

Pruhonice 

May 2002 

vii

Juncaceae Juss., Gen. Pl. 43 (1789) 

Type: Juncus L. 

JUNCACEAE 

Perennial, usually rhizomatous, less often annual herbs, glabrous or with hairy leaf margins; 

rarely leaf abaxial surface with simple, many-celled trichomes. Silica bodies ±absent; 

oxalate raphides absent; stomata paracytic. Stem erect to ascending, rarely procumbent, 

usually terete, leafless or leaf-bearing. Leaves linear or filiform, spirally arranged, rarely 

distichous; sheath closed or open, often auriculate; auricles rarely joined to form a ligule-like 

structure; blades of basal leaves sometimes reduced (cataphylls). Inflorescence terminal, 

rarely pseudolateral (bract resembles a continuation of stem), compound, cymose or 

racemose, panicle-like or anthelate, many-flowered with flowers in many to one terminal 

heads or spike-like clusters; lower inflorescence bracts usually herbaceous; each branch with 

membranous bract and adaxial prophyll; flower bracteoles 1–2 or absent; rarely 

inflorescence reduced to a single terminal or lateral (subterminal) flower. Flowers small, 

usually up to 8 mm long (rarely to 40 mm), actinomorphic, hypogynous, usually 

hermaphrodite, rarely unisexual (dioecious or monoecious). Perianth segments 6, in two 

whorls, glumaceous, usually ±equal, free. Stamens 6 in two whorls or inner whorl reduced; 

filaments filiform to flattened; anthers 2-thecate, 4-sporangiate, oblong to linear, basifixed, 

dehiscence lateral; connective rarely with a projection; pollen in tetrads. Carpels 3, connate; 

ovary superior, unilocular or 3-septate to 3-locular; style 1, distally 3-branched (stigmas), 

papillae ±adaxial. Fruit an orbicular to oblong-ellipsoid loculicidal (rarely circumscissile) 

capsule. Seeds 3–many; endosperm starchy; embryo small, broadly cylindric; outer seed-coat 

hyaline, whitish to brownish, loose to adpressed, sometimes forming apical and/or basal 

appendages; inner seed-coat usually brown to castaneous. 

Seven genera, c. 440 species, almost cosmopolitan, common from temperate to polar regions; 

mostly in the mountains in the tropics. 

Closely related to the South African monotypic Prioniaceae (S.L.Munro & H.P.Linder, Syst. 

Bot. 23: 43–55 (1998)) and to Thurniaceae and Cyperaceae (D.Simpson, in P.J.Rudall et al. 

(eds.), Monocot. Syst. Evol. 497–509 (1995)). 

G.Engelmann, A revision of the North American species of the genus Juncus, with a 

description of new and imperfectly known species, Trans. Acad. Sci. St. Louis 2: 424–458 

(1866), 459–499 (1868); F.G.P.Buchenau, Monographia Juncacearum, Bot. Jahrb. Syst. 12: 

1–495 (1890); F.G.P.Buchenau, Juncaceae, in H.G.A.Engler, Pflanzenr. (iv.36) Heft 25: 

1–284 (1906); F.Vierhapper, Juncaceae, in H.G.A.Engler & K.Prantl, Nat. Pflanzenfam. 2nd 

edn, 15a: 192–224 (1930); M.Barros, Las Juncáceas de la Argentina, Chile y Uruguay, 

Darwiniana 10: 279–460 (1953); D.F.Cutler, Juncales, in C.R.Metcalfe, Anat. Monocot. 4: 

1–358 (1969); L.J.Novara, Contribución al conocimiento de las inflorescencias de Juncus y 

su significación taxonómica, Kurtziana 9: 41–61 (1976); V.S.Novikov, Konspekt sistemy 

roda Juncus L. (Juncaceae), Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 95(5): 111–125 

(1990); S.Snogerup, A revision of Juncus subgen. Juncus (Juncaceae), Willdenowia 23: 

23–73 (1993); H.Balslev, Fl. Neotrop. Monogr. 68: 1–168 (1996); H.Balslev, Juncaceae, in 

K.Kubitzki (ed.), The families and genera of vascular plants, pp. 252–260 (1998); 

R.E.Brooks, S.E.Clemants & J.Coffey Swab, 209. Juncaceae Jussieu. Rush Family, in Flora 

of North America North of Mexico 22: 211–267 (2000); Wu Guofang & S.E.Clemants, 

Juncaceae, in Wu Zhengyi & P.H.Raven, Fl. China 24, Flagellariaceae through 

Marantaceae, pp. 44–69 (2000). 

Throughout the text, unless otherwise stated, the term seed is used for seeds excluding their 

appendages. 

1

2 

SPECIES PLANTARUM — FLORA OF THE WORLD (2002) 

1 Flowers solitary; anthers mucronate (if anthers not mucronate then outer 

tepals at least 15 mm long); auricles never lacerate 

2 Plants not cushion-forming; upper part of stem leafless; flower terminal 

3 Flower bracts 2; lower bract herbaceous, conspicuously longer than 

perianth; upper bract ±equalling perianth; capsule suborbicular to obovoid, 

obtuse, to c. 5 mm long; seeds without conspicuous appendages 1. Rostkovia 

3: Flower bracts 1 or 2, membranous, much shorter than perianth; 

capsule oblong to ellipsoidal, trigonous, acuminate, at least 7 mm 

long; seeds with two distinct appendages 2. Marsippospermum 

2: Plants cushion-forming; upper part of stem densely covered with 

leaves; flower lateral (subterminal, axillary) 

4 Leaves regularly distichous; gynophore developed, elongating during 

capsule ripening 5. Distichia 

4: Leaves ±spirally arranged; gynophore absent 

5 Flower with 1 bracteole; leaf blade with margins finely denticulate, 

with apex acute but not hard nor prickly 4. Patosia 

5: Flowers with 2 bracteoles; leaf blade with smooth margins, with 

apex acute, hard, prickly 3. Oxychloë 

1: Flowers in multi-flowered inflorescences; anthers not mucronate 

(if flowers occasionally solitary then tepals shorter than 10 mm); 

if anthers minutely mucronate then auricles lacerate 

6 Capsule 3-seeded; leaves densely to very sparsely ciliate; flowers always 

with basal bracteoles; leaf sheath closed, rarely partially dissected 6. Luzula 

6: Capsule many-seeded; leaves not hairy; flowers with or without 

bracteoles; leaf sheath open, rarely closed 7. Juncus 

1. ROSTKOVIA 

Rostkovia Desv., J. Bot. (Desvaux) 1: 324 (1809) 

Type: Rostkovia magellanica (Lam.) Hook.f. 

Perennial herbs, erect, glabrous. Stems leafless. Leaves all basal, spirally arranged, linear or 

with reduced blade; stomata restricted to 2 narrow bands of lighter color, one on each side of 

the midrib. Flower bracts 2, with the longer one erect to suberect, much exceeding the 

flower, and the other ±equalling tepals. Flower terminal (pseudolateral), single, large, 

hermaphrodite. Tepals castaneous-brown to dark brown; margins membranous, distally 

involute. Stamens 6; anthers linear; connective with apical mucro to 0.5 mm. Ovary sessile. 

Style with 3 tapering twining stigmas. Capsule subglobose, unilocular or slightly triseptate, 

sessile. Seeds many, ±ovoid; appendage absent; base with a paler patch or apiculate; seedcoat 

with fine longitudinal pattern, with cells narrow elongated and pointed. 

Two species, one endemic to Tristan da Cunha, the other distributed in New Zealand, the 

Sub-Antarctic Islands, the southernmost part of South America to Patagonia, and disjunctly 

in Ecuador. 

Leaves up to 1 mm wide, straight, stiff, acute; style 2.5–4.0 mm 1. R. magellanica 

Leaves usually 1.2–1.7 mm wide, arcuate to circinnate, subacute-subobtuse; 

style 1.5–2.0 mm 2. R. tristanensis


Figure 1. Distichia muscoides. A, habit; B, fruiting shoot showing the capsule lifted on 

gynophore; C, staminate flowering shoot (A, L.Holm-Nielsen et al. 6732, AAU; 

B, B.Øllgaard & H.Balslev 8435, AAU; C, Jameson s.n., K). Rostkovia magellanica. D, habit; 

E, flower; F, fruit. (A-C, B.Øllgaard & H.Balslev 9772, AAU). Scale bars: A, D = 5 cm; B, C, 

E, F = 2 mm. Drawn by K. Tind. Originally published in G.Harling & B.Sparre (eds.), Fl. 

Ecuador 11: 6, fig. 1 (1979), reproduced with permission, from H.Balslev, Fl. Neotrop. 

Monogr. 68: 40, fig. 20 (1996), © 1996, The New York Botanical Garden. 

3

4 


1. Rostkovia magellanica (Lam.) Hook.f., Fl. Antarct. 1: 81 (1844) 

Juncus magellanicus Lam., Encycl. 3: 266 (1789); Rostkovia sphaerocarpa Desv., J. Bot. (Desvaux) 1: 327, pl. 

12, fig. 2 (1809), nom. illeg. T: Chile. Magallanes: Magellan Strait, P.Commerson s.n.; holo: P, photo 

AAU; iso: P, US. 

Rostkovia magellanica (Lam.) Hook.f. f. pumila Albov, Revista Mus. La Plata, Secc. Bot. 7: 353 (1896). 

T: Argentina, Tierra del Fuego, left bank of Río Grande, 6 Mar 1896, N.Albov 808; holo: LP, photo NY; iso: SI. 

Illustrations: F.G.P.Buchenau, in H.G.A.Engler, Pflanzenr. (iv.36) 25: 40, fig. 34 (1906); M.Barros, 

Darwiniana 10: 306, fig. 8 (1953); H.Balslev, Brittonia 31: 244, fig. 1 (1979); H.Balslev, Fl. Neotrop. 

Monogr. 68: 40, fig. 20, D-F (1996). 

Map: H.Balslev, Brittonia 31: 245, fig. 2 (1979). 

Perennials, loosely caespitose, 15–30 cm tall; rhizome erect or ascending, sometimes short 

creeping, 1.5–2.5 mm diam., usually covered with brown 1.0–1.5 cm long scales inserted 

0.5–1.0 cm apart. Stem 0.5–1.0 mm diam.; cataphylls 2–4 per stem, to 7 cm long, longitudinally 

ridged, light brown, with smooth red-brown margins; rudimentary blade to 4 mm 

long. Basal leaves 15–25 cm long, to 1 mm wide; tip acute; auricles c. 1 mm long, rounded; 

blades straight, stiff, ±flat, slightly trigonous or canaliculate, narrowly grooved along abaxial 

midrib. Flower bracts 2; lower bract long, herbaceous, 4 cm long; upper bract scale-like, to 

1 cm long. Pedicel 1–3 mm long. Tepals equal, lanceolate, 6–7 mm long, dark reddish 

brown, gradually scarious toward the margin, obtuse, seemingly acuminate because of the 

involute scarious margins. Stamens c. 4.5 mm long; anthers c. 2.5–3.0 mm long, mucronate; 

filaments linear, flat, to 3 mm long, usually shorter than anthers; style 3.0–4.5 mm long; 

stigmas c. 2.5–3.0 mm long. Capsule c. 4 mm in diam., thick-walled, dark brown to black, 

shining. Seeds oblong to ovoid, c. 1.2–1.7 × 0.7–0.9 mm, dark, almost blackish, glossy with 

fine color pattern of longitudinal blackish rectangles. 2n=56, fide E.J.Beuzenberg & 

J.B.Hair, New Zealand J. Bot. 21: 15 (1983). Fig. 1. 

Rostkovia magellanica occupies a wide and very disjunct range: New Zealand main islands 

and Antipodean Is., South Georgia, Falkland Islands, southernmost Chile and Argentina, and 

Ecuador. 51: ATP, NZS. 83: ECU. 85: AGS, CLS. 90: FAL, SGE. Mossy bogs and wet 

depressions, from sea level (in the south) up to 3400 m (in Ecuador). Map 1. 

51. ANTIPODEAN IS.: Campbell Is., Mt Lyall, 19 Dec 1975, D.R.Given 9173 (K); Campbell Is., Dec 1840, 

J.D.Hooker 640 (K). 83. ECUADOR: Morona–Santiago, between Alao and Huamboya near pass, 3550–3950 m, 

B.Øllgaard et al. 38239 (AAU, F, MO, NY, QCA); S.Laegaard 55422 (AAU, QCA, W). 85. ARGENTINA 

SOUTH: Argentina, Patagonia, Río Fosiles, N of Lago San Martín, 1000 m, 6 Jan 1909, C.J.F.Skottsberg s.n. 

(UPS); Santa Cruz, Lago Argentino, Brazo Onelli, 660 m, F.B.Vervoorst 4632 (K, LIL); Tierra del Fuego, 

Sierra Alvear, Paso Garibaldi, 800 m, D.M.Moore 1587 (K). CHILE SOUTH: Tierra del Fuego, Estancia 

Camerón, Campo Zapata, 21 Mar 1964, D.M.Moore 1114 (K); Tierra del Fuego west, Fjordo Agostini, 22 Feb 

1929, H.Roivainen s.n. (H). 90. FALKLAND IS.: Port Stanley, Mt Williams, 28 Feb 1904, S.Birger (LD). 

SOUTH GEORGIA: Cumberland Bay, Darthmouth Pt., 2 Jan 1959, W.Bonner 95 (BM); King Edward Point, 

28 Feb 1957, Smith 1074 (K). 

2. Rostkovia tristanensis Christoph., Results Norweg. Sci. Exped. Tristan da Cunha 

11: 3 (1944) 

T: Tristan da Cunha, above Soggy Plain, 1100–1200 m, 19 Feb 1938, Y.Mejland 1516; holo: O, photo NY; iso: 

BM, GH, K, LD, O, S. 

Illustrations: E.Christophersen, Res. Norweg. Sci. Exped. Tristan da Cunha 11: 4 and 5, figs. 2, 3 (1944). 

Perennials, loosely caespitose, to 16 cm tall; rhizome creeping to ascending, often 

stoloniferous, covered with scales; stems erect, rigid, 0.7–1.0 mm in diam.; cataphylls 

coriaceous, with two prominent veins, to 3–5 cm long; rudimentary blade to c. 2 mm; basal 

leaves usually to 10–12 cm long, 1.2–1.7 mm wide, flat, canaliculate, arcuate to circinnate, 

subobtuse to subacute. Flower bracts 2; lower bract 1.2–1.9 (–2.5) cm long; upper bract 

slightly shorter to ±equalling perianth. Pedicel short. Tepals subequal, lanceolate, 

castaneous-brown; margins membranous-stramineous, ±involute; outer tepals 8–10 mm long; 

inner tepals7–9 mm long. Stamens 5–6 mm; anthers c. 2 mm long; filaments to 4 mm long; 

style 1.5–2.0 mm; stigmas 1.5–2.5 mm. Capsule obovoid, 4.5–5.5 mm long, ±brown, shining 

to dull. Seeds narrowly ovoid, 1.0–1.2 mm long, shining, with fine longitudinal pattern.

JUNCACEAE (Rostokovia) 

Endemic to Tristan da Cunha. Known only from the rich type collection. 90: TDC Tristan da 

Cunha, above Soggy Plain, 19 Feb 1938, Y.Mejland 1516 (BM, GH, K, LD, photo NY, O, S). 

Mountain Empetrum heathland at 1100–1200 m. Map 2. 

On Gough Island (90. TDC), plants of intermediate character between R. magellanica and 

R. tristanensis were collected in 1956 [Hog's Tooth, c. 700 m, peat among trachyte blocks, 

N.M.Wace 132 (BM)]. They approach the former in having narrow, straight, erect, stiff 

leaves and shorter tepals; they are close to R. tristanensis in short anthers (c. 1.7–1.9 mm), 

shorter than filaments, probably short stigmas and paler perianth colour. As both 

R. tristanensis and the Gough Is. plants are known from rather limited material, resolution of 

this discrepancy must await fuller collections. Possibly, R. tristanensis may vary in its leaf 

characters. 

2. MARSIPPOSPERMUM 

Marsippospermum Desv., J. Bot. (Desvaux) 1: 328 (1808); Juncus sect. Marsippospermum 

(Desv.) E.Mey., Syn. Junc. 52 (1822) 

Type: Marsippospermum calyculatum Desv., nom. Illeg., = Juncus grandiflorus L.f. 

Perennial herbs, erect, glabrous. Leaves 1–2 (–3) cauline; sterile shoots with 1–3 (–5) 

cauliform terete leaves; pith parenchymatous; cataphylls distinctly developed; stomatal lines 

distributed evenly. Flower bracts 1–2, short, membranous. Flowers borne singly, terminal, 

large, usually 10–40 mm long; tepals linear-lanceolate, acuminate, pale. Stamens 6; anthers 

linear, longer than filaments; connective with or without apical projection. Ovary sessile. 

Style with 3 ±tapering stigmas. Capsule oblong-trigonous or ellipsoid-trigonous, acuminate, 

imperfectly triseptate, with upper part recurved after dehiscence. Seeds many; seed-coat 

forming two long tail-like appendages. 

Four species; one endemic to New Zealand, the others confined to southernmost South 

America and Falkland Islands. 

1 Connective with a distinct apical projection at least 0.7 mm long; overall 

length of seeds less than 2.7 mm 

2 Connective with an apical projection 1.8–2.0 mm long; outer tepals 

1.1–1.5 cm long; leaves of sterile shoots c. 0.5 mm in diam. 

[South America] 4. M. reichei 

2: Connective with an apical projection 0.7–0.9 mm long; outer tepals 

1.7–2.7 cm long; leaves of sterile shoots 0.6–0.9 mm in diam. 

[New Zealand] 3. M. gracile 

1: Connective ±without apical projection; overall length of seeds 

at least 4.0 mm 

3 Outer tepals usually 3.2–4.0 cm long; anthers usually 8–10 mm long; 

capsule 1.7–2.5 cm long; seeds 5.0–6.5 mm long including appendages 1. M. grandiflorum 

3: Outer tepals usually 1.6–2.3 cm long; anthers usually 4–7 mm long; 

capsule 1.1–1.5 cm long; seeds 4.0–5.0 mm long including appendages 2. M. philippii 

1. Marsippospermum grandiflorum (L.f.) Hook., Icon. Pl. new ser., 2 [6]: tab. 533 

(1843) 

Juncus grandiflorus L.f., Suppl. Pl. 1: 209 (1781); Marsippospermum calyculatum Desv., J. Bot. (Desvaux) 

1: 330 (1808), nom. illeg.; Rostkovia grandiflora (L.f.) Hook.f., Fl. Antarct. 2: 357 (1847). T: Tierra del 

Fuego, Baeck; holo: S; plants from the same gathering: 1774, J.R.Forster, & G.Forster & Sparmann; possible 

iso: C, fide B.Hansen & P.Wagner, Allertonia 7: 337 (1998). 

Illustrations: W.J.Hooker, Icon. Pl. new ser., 2 [6]: tab. 533 (1843); M.Barros, Darwiniana 10: 310, fig. 9 

(1953); S.Parkinson, in D.M.Moore, Fl. Tierra del Fuego 320, fig. 254, 322, fig. 260 (1993). 

Map: D.M.Moore, Fl. Tierra del Fuego 318 map 36 j (1993). 

5

6 


Perennials, loosely caespitose; rhizome stout, creeping, with internodes short, to 1–2 cm 

long; stems in rows, usually 25–40 (–60) cm long, c. 1.5 mm in diam. Cataphylls 2 (–3), 

loose, pale brown; sheaths obtuse; blade setiform, usually 2–4 mm; cauline leaf usually 

single; auricles short, obtuse. Leaves of sterile shoots single, cauliform, terete, equalling to 

much exceeding stems, c. 2 mm in diam., ±smooth to slightly striate; tip acute to subobtuse. 

Bracteoles 2, membranous, broadly ovate, obtuse, usually 2–4 mm long. Tepals linearlanceolate, 

pale stramineous to scarious, irregularly unequal; outer tepals usually 3.2–4.0 cm 

long; inner tepals c. 2.0–2.5 (–3.5) cm long. Stamens 6, much shorter than perianth, usually 

10–11 mm long. Anthers 8–10 mm long; pollen sacs slightly overtopping connective; tips 

later erecto-patent and hence anthers bicorniculate at apex. Filaments to c. 2 mm long. Style 

c. 5–6 mm long; stigmas 8–10 mm long. Capsule oblong-ellipsoidal, ±trigonous, acuminate, 

usually 1.7–2.2 (–2.5) cm long, pale brown, shining. Seeds pale brown, ±ellipsoidal, with 

body c. 1.0 mm long, 0.4–0.5 mm wide, or 5.0–6.5 mm long including the two whitish 

subequal tail-like appendages each 1.7–2.4 mm long. Fig. 2. 

Southern South America, from Los Lagos, Chile, and Neuquén, Argentina, to Tierra del 

Fuego, and Falkland Is. 85: AGS, CLS. 90: FAL. Bogs, peaty grasslands, wet sands, 

moorland, usually at altitudes below 800 m. Map 3. 

85. ARGENTINA SOUTH: Santa Cruz, Lago Argentino, Brazo Onelli, 700 m, F.B.Vervoorst 4602 (W); Tierra 

del Fuego, Isla de los Estados, Bahía Colnett, 28 Feb 1968, D.M.Moore 2043 (UPS); SW Patagonia, Cerro 

Buenos Aires, 900 ft, 4 Jan 1959, P.W.James 681 (BM). CHILE SOUTH: Hermite Is., Cape Horn, 

J.D.Hooker (K); Taitao Peninsula, San Rafael, between 46° and 47°, M.Gusinde 476 (W); Tierra del Fuego, 

Bahía Blanca, D.M.Moore 2281 (H, K); Tierra del Fuego, Porvenir, P.Dusén 261 (UPS); Los Lagos, Chiloé, 

Cordillera San Pedro, 8 Oct 1958, Godley 418c (BM). 90. FALKLAND IS.: W Falkland, Mt Cook, 800', 

Vallentin s.n. (K); Stanley Harbour, Klondyke, 14 Nov 1907, C.J.F.Skottsberg 22 (LD, UPS) 

2. Marsippospermum philippii (Buchenau) Hauman, Anales Mus. Nac. Hist. Nat. 

Buenos Aires 27: 295 (1915) 

Marsippospermum grandiflorum var. philippii Buchenau, Bot. Jahrb. Syst. 12: 68 (1890); Rostkovia gracilis 

Phil., Linnaea 29: 75 (1857), nom. illeg. T: Chile, ‘In monte Doce del Febrero inter lacus Todos los Santos 

& Nahuelhuapi’, 1600 m, Fonk; holo: SGO, n.v. 

Illustration: M.Barros, Darwiniana 10: 310, fig. 9 (1953). 

Perennials, densely caespitose; rhizome stout, branched, ascending or short with internodes 

very abbreviated; stem 15–25 (–33) cm long, c. 1 mm in diam.; cataphylls 3–4 (–5), obtuse, 

pale brown with blade mucroniform and 0.3–1.5 mm long; cauline leaf usually single; 

auricles narrow, distinct; sterile shoots usually single-leaved. Leaves overtopping stem; 

apex acute, smooth. Bracteoles 2, membranous, obtuse, broadly rounded, c. 1.0–1.5 mm 

long. Tepals linear-lanceolate, acuminate, scarious to stramineous, unequal; outer tepals 

1.6–2.3 cm long; inner tepals c. 1.3–1.5 cm. Stamens 6; anthers 4–7 mm long; filaments 

(0.7–) 1.2–1.5 mm long; ovary c. 3 mm long; style c. 3 mm long; stigmas c. 5 mm long. 

Capsule ellipsoid-trigonous, acuminate, brown, shining, 1.1–1.5 cm long. Seeds 1.0–1.1 mm 

long, or 4.0–5.0 mm long including two whitish subequal tail-like appendages 1.2–1.5 mm 

and 1.5-1.7 mm long. 

Confined to the southern part of C Chile and Neuquén and Río Negro in SW Argentina. 

85: AGS, CLC. Peaty grassland, bogs, up to 2200 m. Map 4. 

85. ARGENTINA SOUTH: Río Negro, Cerro López, 1700 m, A.Burkart (SI 6177). CHILE CENTRAL: 

Baños de Chillán, Aguas Calientes, 2200 m, Mar 1927, E.Werdermann 1302 (K, S). 

3. Marsippospermum gracile (Hook.f.) Buchenau, Junc. S.-Amer. 374 (1879) 

Rostkovia gracilis Hook.f., Fl. Antarct. 2(1): 83, tab. 47 (1844); Juncus gracilis (Hook.f.) Walp., Ann. 

Bot. Syst. 1: 878 (1849), nom. illeg., non R.Br.; Juncus hookeridis Steud., Syn. Pl. Glumac. 2: 307 (1855). 

T: New Zealand, Antipodean Is., Lord Auckland’s Islands, above Rendezvous Harbour, Nov 1840, 

J.D.Hooker; lecto: K, fide E.Edgar in L.B.Moore & E.Edgar, Fl. New Zealand 2: 58 (1970).

JUNCACEAE (Marsippospermum) 

Figure 2. Marsippospermum grandiflorum. A, habit; B, anther; C, flower; D, seed. 

M. reichei. H, habit; E, flower; F, stamen. M. gracile. G, flower. (A–D, D.M.Moore 2043, 

UPS; C.J.F.Skottsberg 22, UPS; H, C.J.F.Skottsberg 725, S, UPS; E, F, P.Dusén 5905, UPS; 

G, E.Hörandl & Hadaček 8089, W). Scale bars: A, H = 8 cm; B, F = 3 mm; C, E, 

G = 2 mm; D = 2 mm. Drawn by © E.Smrčinová. 

7

8 


Rostkovia novae-zelandiae Buchanan, Trans. & Proc. New Zealand Inst. 4: 227 (1872); Marsippospermum 

gracile var. novae-zelandiae (Buchanan) Buchenau, Bot. Jahrb. Syst. 12: 69 (1890). T: New Zealand. 

Nelson Mtns, H.H.Travers; syn: K; ‘mountains of Canterbury’ J.F.J. von Haast; syn: K. 

Illustrations: J.D.Hooker, Fl. Antarct. 2(1): 83, tab. 47 (1844); F.G.P.Buchenau, in H.G.A.Engler, Pflanzenr. 

(iv.36) 25: 35, fig. 28 (1906); L.B.Moore & J.B.Irwin, Oxford Book New Zealand Pl. 199, fig. 2 (1978). 

Perennials, densely caespitose; rhizome horizontal, rarely ascending, with internodes very 

abbreviated; stems crowded in dense rows, usually 10–30 cm tall; lower cataphylls ovate to 

rounded, greyish stramineous, wrinkled, with blade reduced to mucro 0.3–0.7 mm long; 

upper cataphylls usually up to 5–6 cm long, obtuse above, straw-brown, rarely reddish 

brown, slightly shining to dull, with blade fragile, subulate, usually to 1.5 cm long; shoots 

erect, extravaginal. Leaves terete, much exceeding stems, ±rigid, ±striate, bright green, 

±shining above, usually 20–35 cm long, 0.6–0.9 mm in diam., acute, ±papillose near apex; 

basal leaves 1–3 per shoot; cauline leaves usually single, with long sheath. Bract(eole) 

solitary, ovate, obtuse, membranous, 2.0–2.5 mm long. Tepals unequal, erect, entire, 

acuminate to subaristate, straw-brown to stramineous, margins ±membranous; outer tepals 

usually 1.7–2.7 mm long, inner tepals usually 1.3–1.9 mm long, up to 1.5 mm wide. Stamens 6; 

anthers c. 5 mm long, apiculate; filaments c. 1.7–2.0 mm long; connective prominent; mucro 

0.7–0.9 mm long; style dark, c. 2.5 mm; stigmas dark, c. 5 mm. Capsule oblong-ellipsoidal, 

subacuminate, mucronate, usually 8–10 mm long, pale brown to castaneous brown. Seeds 

straw-brown, oblong ellipsoidal, c. 0.9–1.1 × c. 0.6 mm; seed-coat distinct; overall length 

2.3–2.7 mm including unequal appendages c. 0.4–0.6 mm and c. 0.8–1.1 mm long. Fig. 2. 

Endemic to South Island and Antipodean Islands, New Zealand. 51: NZS, ATP. Moist places 

among rocks, marshes, moist alpine meadows, 150–2100 m. Map 5. 

51. NEW ZEALAND SOUTH: South Is., Fiordland Nat. Park, from Homer Hut to Gertrude Valley, 

1100–1200 m, E.Hörandl & Hadaček 8089 (W); Canterbury, Arthur Pass, 1874, S.Berggren (K); Canterbury, 

Arthur Pass, Blimit Peak, 20 Jan 1928, W.R.B.Oliver (K); Canterbury, Kelly’s Hill, Feb 1874, S.Berggren (LD, 

S, UPS); Nelson, Mt Arthur, T.F.Cheeseman s.n. (H). ANTIPODEAN IS.: Campbell Is., Mt Azimuth, 

D.R.Given 9115 (K). 

4. Marsippospermum reichei Buchenau, Ber. Deutsch. Bot. Ges. 19: 160 (1901) 

Rostkovia reichei (Buchenau) Hosseus, Trab. Inst. Bot. Farmacol. 33: 24 (1915), n.v. T: Chile, 'In regione 

nivis aeternae (1000 m s.m.) montis Patagoniae australis in oriente urbis Bahia Ultima Esperanza, c. 50°30' ', 

Feb 1900, K.F.Reiche; syn: n.v. 


Map: D.M.Moore, Fl. Tierra del Fuego 324, map 37, a (1993). 

Perennials, densely caespitose; rhizome short, ascending, with internodes very abbreviated; 

stems usually 5–10 cm tall, to 0.7 mm in diam. Cataphylls castaneous brown at base, shining, 

±gradually tapered to ±obtuse apex; blade setiform, 2–5 mm long. Cauline leaves 1–3, subbasal; 

sheaths to 3–4 cm long; blade to c. 5–7 cm long. Sterile shoots (2–) 3 (–4)-phyllous; 

leaves filiform, c. 0.5 mm in diam., usually 10–15 (–20) cm long, much overtopping stem, 

with tip acute. Bracteole single, membranous, obtuse, to c. 1 mm long. Tepals linear to 

linear-lanceolate, c. 0.5 mm wide, acuminate, membranous to stramineous, unequal; outer 

tepals usually (1.1–) 1.3–1.5 cm long; inner tepals (0.9–) 1.1–1.2 cm long. Stamens 6, 5–6 mm 

long; anthers usually 3.0–3.5 mm long; connective with distinct 1.8–2.0 mm long apical 

projection; filaments 1.1–1.5 mm long. Style short, c. 1.2–1.5 mm long; stigmas c. 4 mm 

long. Capsule oblong-trigonous, acuminate, usually 7–9 mm long, pale brown, shining. Seeds 

castaneous-brown, 0.8–0.9 mm long, or 1.7–2.0 mm long including two unequal whitish 

appendages c. 0.3 and c. 0.5 mm long. Fig. 2. 

Southernmost South America, from Neuquén southwards to Tierra del Fuego in Argentina, 

province of Magallanes and Islas Wollaston in Chile. 85: AGS, CLS. Wet rocks, gravelly 

open substrates along streams, up to 1000 m. Map 6. 

85. ARGENTINA: Santa Cruz, Lago San Martín, Río Fossiles, 1000 m, 27 Mar 1905, P.Dusén s.n. (K); 

loc. cit. P.Dusén 5905 (UPS); loc. cit. P.Dusén 13223 (SI). CHILE/ARGENTINA: S Patagonia, Sierra 

Baguales, 1500 m, C.J.F.Skottsberg 725 (S, UPS).


3. OXYCHLOË 

Oxychloë Phil., Fl. Atacam. 52 (1860); Distichia sect. Oxychloë (Phil.) Benth. & Hook.f., 

Gen. Pl. 3: 867 (1883). 

Type: Oxychloë andina Phil. 

Andesia Hauman, Anales Mus. Nac. Hist. Nat. Buenos Aires 27: 287 (1915). T: Andesia bisexualis (Kuntze) 

Hauman 

Perennial herbs, cushion-forming, glabrous; cushions dense, prickly. Rhizome branched, 

ascending. Stems ±erect, ±regularly forked, with proximal part densely covered with 

persisting wilted leaf bases. Leaves regularly and densely spirally inserted along stem; blade 

equal to or longer than sheath, terete, smooth, with apex acute, hard-pointed, prickly. 

Inflorescence reduced to a single lateral (axillary) subterminal flower with two scarious 

bracteoles; pedicel of variable length. Flowers unisexual or hermaphrodite (plants 

polygamous-dioecious or diclino-dioecious). Tepals glumaceous or papery with margins 

hyaline, lanceolate or ovate-lanceolate, often unequal. Staminate flower with 6 stamens 

shorter than or equalling tepals; anthers elongated, mucronate, much longer than the very 

short flattened ±triangular filament. Stigmas tapering. Capsule trilocular to triseptate, 

loculicidal to circumscissile. Seeds many, ovoid to pyriform; seed-coat distinct, forming 

inconspicuous appendage, whitish-yellowish, minutely reticulate, with cells slightly 

elongated longitudinally. 

Five species at high altitudes of the Andes in South America, from Peru to north Patagonia in 

Argentina. The overall similarity among the species is high, the species are difficult to 

cultivate, and the genus requires a thorough taxonomic and population study. 

1 Leaf blade 40–60 mm long 5. O. mendocina 

1: Leaf blade 10–30 mm long 

2 Leaf blade up to 20 mm long 

3 Perianth c. 7–8 mm long; capsule 4–5 mm long, shorter than perianth; 

all flowers unisexual, with pistillate and staminate flowers on 

different shoots 3. O. castellanosii 

3: Perianth to 5 mm long; capsule 8–10 mm long, exceeding perianth; 

hermaphrodite and unisexual flowers on different shoots 4. O. haumaniana 

2: Leaf blade 20–30 mm long 

4 Capsule 8–10 mm long, longer than perianth; seeds 1.0–1.3 mm long; 

all flowers unisexual, with pistillate and staminate flowers on 

different shoots 1. O. andina 

4: Capsule c. 6 mm long, ±equalling perianth; seeds 0.6–0.8 mm long; 

hermaphrodite and unisexual flowers on different shoots 2. O. bisexualis 

1. Oxychloë andina Phil., Fl. Atacam. 52, tab. 6c (1860) 

Distichia andina (Phil.) Benth. & Hook.f., Gen. Pl. 3: 867 (1883). T: Chile, Antofagasta, Atacama desert, 

Zorras, 3200 m, Feb 1854, R.A.Philippi s.n.; lecto: SGO63055, fide H.Balslev, Fl. Neotrop. Monogr. 68: 49 

(1996); isolecto: K, SGO37716. 

Distichia macrocarpa Wedd. ex Buchenau, Abh. Naturwiss. Vereine Bremen 4: 123 (1874). T: Bolivia, La 

Paz, near Sorata, Vancuiri near Chuchu, 4500–5000 m, Jan–Mar 1857, G.Mandon 1442; holo: GOET, n.v.; iso: 

G, GH, K, NY, P, S. 

Illustrations: F.G.P.Buchenau, Junc. S.-Amer., Plate 3 (1879); M.Barros, Darwiniana 10: 297, fig. 4 (1953). 

Perennials, compact, cushion-forming. Stems erect, much branched, distal part with leaves 

8–10 cm. Leaves rigid and conical; blade 20–30 mm long, to 1.2 mm wide; auricles 

membranous, rounded, sometimes forming a ligule-like adaxial structure. Bracteoles 

scarious, unequal, 3.0–4.5 mm long. Flowers unisexual; pedicels 1–3 cm long, ±equal in both 

sexes. Staminate flowers with tepals narrowly lanceolate, scarious, stramineous, rarely 

castaneous; outer tepals 6.5–7.5 mm long; inner tepals 7.5–8.0 mm long; stamens 6; anthers 

9

10 


4–5 mm long, mucronate; filaments 0.3–0.5 mm long; pistil rudimentary. Pistillate flowers 

slightly smaller; tepals broadly lanceolate, 6–8 mm long; ovary ellipsoid, 2 mm long; style 

c. 2 mm long, stigmas filiform, elongated, c. 2–4 mm long, exserted. Capsule ovoid, with 

rigid, shining wall, 8–11 mm long, 4.0–4.5 mm wide. Seeds 1.0–1.3 mm long, 0.8–1.0 mm 

wide. 2n=16 (Sasaki, 1937, n.v., fide H.Balslev, Fl. Neotrop. Monogr. 68: 32 (1996) 1996: 

32; an old, doubtful count). Fig. 3. 

Confined to the high Andes of Peru, Bolivia, N Chile and NW Argentina: Salta to San Juan. 

83. PER, BOL. 85. AGW, CLN. Humid bogs, usually above 4000 m. Map 7. 

83. PERU: Puno, San Antonio de Esquilache, 4700 m, D.Stafford 736 (F, K, WIS). BOLIVIA: La Paz, 

Murillo, Pacocota Pampa, 4200 m, E.Asplund 190 (UPS); Potosí, Sud Lípez, Cerro Tapaquillcha, 4200 m, 

M.Liberman 456 (LPB). 85. ARGENTINA NORTHWEST: Catamarca, Tinogasta, Tres Quebrados, 4200 m, 

F.B.Vervoorst 3234 (LIL, W); La Rioja, Cordillera de la Boca – Carneritos, A.Krapovickas & A.T.Hunziker 

5671 (CORD); Salta, Valle La Ollada, between Cerro La Apacheta and the foot of Nevado de Cachi, 

4500–4750 m, V.Núñez 581 (MCNS); San Juan, Valle del Cura, R.Pérez Moreau 30/45 (WU); San Juan, 

Reserva San Guillermo, Vega de los Corrales, E.Nicora, E.R.Guaglianone & A.M.Ragonese 8317 (SI, MCNS). 

CHILE NORTH: Ríofrío, 24°50’ S, 3350 m, R.A.Philippi (SGO); Atacama, Chanaval, 7 km NE of Cerro 

Valiente, 3400 m, I.M.Johnston 4825 (F, GH, K, P, S, US). 

2. Oxychloë bisexualis Kuntze, Revis. Gen. Pl. 3(2): 321 (1898) 

Andesia bisexualis (Kuntze) Hauman, Anales Mus. Nac. Hist. Nat. Buenos Aires 27: 290 (1915). T: Argentina, 

Paso Cruz, 34° N, 2800 m, O.Kuntze; lecto: K, designated here; photo S. 

Illustrations: L.Hauman, Anales Mus. Nac. Hist. Nat. Buenos Aires 27: 291, plate vi, fig. 1c (1915); M.Barros, 

Darwiniana 10: 306, fig. 8 (1953). 

Dense perennials, cushion-forming. Stems ±erect, much-branched. Leaves only in the most 

distal part; sheath c. 3–4 cm long; blade rigid, semicylindrical in T.S., canaliculate at base, 

2–3 cm long, 1.5–2.0 mm wide; auricles membranous, fused, forming a ligule-like structure. 

Bracteoles membranous-scarious, unequal; lower bracteoles 4.5 mm long; upper bracteoles 

4 mm long. Flowers hermaphrodite or unisexual (plants polygamo-dioecious); pedicel thin, 

to 5 cm long, exceeding leaves. Tepals 6.5–7.5 mm long. Stamens 6; anthers elongated, 

3.5–4.0 mm long, mucronate; filaments 0.3–0.5 mm long; ovary ovoid, brownish, 2.5–3.0 mm 

long; style slender, 3.5–4.0 mm; stigmas exserted, 1.5–2.0 mm. Capsule ovoid, ±trigonous, 

brownish, shining, c. 6 mm long, 2.0–2.5 mm wide. Seed minutely reticulate, c. 0.6–0.8 mm 

long. 

Endemic to the Andean region in Argentina, from N Patagonia to Salta, and in N Chile. 85: 

AGS, AGW, CLN. Wet bogs, springs, 3300 to 4400 m. Map 8. 

85. ARGENTINA NORTHWEST: Salta, 15 km N of Abra del Gallo, 10 km S of Mina La Poma, 4400 m, 

L.Novara, A.Charpin & L.Albornoz 5723 (MCNS); A.Charpin & L.Novara 20771 (G, MCNS); San Juan, Vega 

de Santa Rosa, 3300 m, A.T.Hunziker & O.Caso 4804 (CORD); Mendoza, Río Blanco, L.Hauman (BA 15631). 

3. Oxychloë castellanosii Barros, Lilloa 23: 417, fig. 2 (1950) 

T: Argentina, San Juan, Cordillera de Colangil, ‘Los Champones’, 5–6 Feb 1950, A.Castellanos s.n.; syn: 

LIL16042, LIL16069, LIL16133. 

Oxychloë simulans Barros, Darwiniana 10: 301 (1953). T: Argentina, La Rioja, Sarmiento, Cordillera de 

los Andes ‘El Zanjón’, 3500 m, 6 Feb 1949, A.Krapovickas & A.T.Hunziker; holo: BAB5817. 

Illustrations: M.Barros, Darwiniana 10: 300, fig. 5, 6 (1953). 

Dense perennials, cushion-forming. Stems erect, branched, (6–) 8–12 cm long; base covered 

with old sheaths; distal part with leaves usually not longer than 2–5 cm. Leaves imperfectly 

distichous; sheaths 1.2–1.5 cm long; auricles broad, obtuse, ±membranous; blades erectopatent, 

rigid, canaliculate at base, cylindrical and gradually narrowing into pungent tip, 

10–15 (–20) mm long, 0.7–1.2 mm wide. Bracteoles membranous-scarious, ovate, ±obtuse, 

unequal, 3.0–4.5 mm long. Flowers all unisexual. Staminate flowers on pedicels 1.2–2.0 cm; 

tepals scarious-membranous, with base stramineous to subcoriaceous; outer tepals 6.5–7.0 mm 

long; inner tepals 7–8 mm long; stamens 6; anthers 4.0–4.5 (–5.5) mm long, mucronulate, with 

mucro c. 0.3 mm long; filaments 0.5 mm long. Pistillate flowers on pedicels 5–7 (–15) mm

JUNCACEAE (Oxychloë) 

Figure 3. Oxychloë andina. A, habit; B, leaf; C, staminate flower, note two bracteoles at 

base of flower; D, stamen; E, pistillate flower at anthesis with exposed style; F, gynoecium; 

G, pistillate flower with mature fruit and remaining style and stigmas. (A, B, F, E, H.Balslev 

1150, NY; G, Werdermann 937, C; D, H.Balslev 1147, NY). Scale bars: A = 5 cm; B–G = 1 mm. 

Drawn by H. Fukuda. Reproduced with permission, from H.Balslev, Fl. Neotrop. Monogr. 

68: 50, fig. 24 (1996), © 1996, The New York Botanical Garden. 

11

12 


long, slightly smaller than male; tepals 6.5–7.0 mm long, more broadly lanceolate; ovary 

turbinate; style c. 3 mm long; stigmas not exserted or slightly so, 1.0–2.5 mm long. Capsule 

unilocular, ellipsoid to oblong-obovoid, rounded at the apex, 4.0–5.0 mm long. Seed 

ellipsoidal, c. 0.8–1.0 mm long. 

Endemic to the west of Argentina, in the Cuyo Region, San Juan Province, and La Rioja 

Province. 85: AGW. Humid bogs and slopes, wet plains and stream banks in high elevations, 

above 4000 m. Map 9. 

85. ARGENTINA NORTHWEST: San Juan, Cuyo, Cordillera de Colanguil, Los Champones, A.Castellanos 

s.n. (W, isosyn., duplicates of LIL16042 & 16133); San Juan, Reserva San Guillermo, Mina Los Caranchos, 

4000 m, Pujalte 103 (SI). 

4. Oxychloë haumaniana (Barros) Barros, Darwiniana 10: 303 (1953) 

Andesia haumaniana Barros, Lilloa 23: 419 (1950). T: Argentina, San Juan, Río Cura, 3900 m, 16 Feb 

1950, A.Castellanos; holo: LIL16136. 

Illustrations: M.Barros, Darwiniana 10: 304, fig. 7 (1953); M.Barros, Lilloa 23: 419, fig. 3 (1950). 

Dense perennials, cushion-forming, 5–10 cm tall. Stem erect, much branched, covered with 

old sheaths, with living leaves only in distal part. Leaves imbricate, imperfectly distichous; 

sheaths brownish, 10–50 mm long, 4–7 mm wide; auricles asymmetrical, ligulate, 

membranous; blade narrowly cylindrical, rigid, surface minutely striate, 10–15 mm long, 

1.0–1.2 mm wide; tip prickly. Bracteoles unequal; lower bracteoles subacute, 4–5 mm long, 

shortly mucronate; upper bracteoles subobtuse, c. 2.5 mm long. Flowers hermaphrodite or 

unisexual; perfect flowers on short peduncle, scarcely overtopping leaves; tepals acute, 

narrowly lanceolate, brownish-purplish with scarious margins, subequal, c. 5 mm long. 

Stamens 6; anthers c. 2.5 mm long, mucronulate; filaments 0.5–1.0 mm long. Unisexual 

flowers similar to the perfect ones, lacking one of the fertile whorls. Capsule hard-walled, 

ovoid, brownish, 8–10 mm long. Seeds obovoid, reticulate, whitish, 0.7–0.9 mm long. 

Rare, endemic to San Juan Province in Argentina; known from the type collection only. 

85: AGW. Wet slopes and plains in high Andes, at 3900 m. Map 10. 

Similar to O. bisexualis but differs in having shorter leaf blade, up to 15 mm long, and 

smaller flowers, usually less than 5 mm. 

5. Oxychloë mendocina Barros, Lilloa 28: 282, fig. 2 (1957) 

T: Argentina, Mendoza, Tristeza, China Muerta, 23 Feb 1953, A.Castellanos; holo: LIL15365. 


Dense perennials, compact, cushion-forming, usually to 8–10 cm tall. Stem erect, much 

branched, with basal internodes covered with old sheaths and fresh leaves only in the most 

distal part. Sheaths with membranous margins, 3–5 cm long; auricles forming a bilobed 

ligule; blade cylindrical, tapered into pugent apex, 4–6 cm long, 1–2 mm wide. Bracteoles 

unequal, ovate, membranous; lower bracteoles 4.5–5.0 mm long. Flowers hermaphrodite or 

unisexual, with the former on long filiform peduncules 3–5 cm long. Tepals lanceolate, 

purplish-castaneous, with margins membranous, subequal; outer tepals 7 mm long; inner 

tepals 7.5–8.5 mm long. Stamens 6; anthers mucronulate, 4.0–4.5 mm long; filaments 

0.3–0.5 mm long. Female flowers with filiform staminodes; ovary turbinate; style c. 2 mm 

long, large; stigma exserted, c. 4–5 mm long. Capsule ovoid to ellipsoidal, 7.0–7.5 mm long, 

shorter than perianth. Seeds ellipsoidal, 0.8–0.9 mm long. 

Endemic to Mendoza Province, W Argentina. 85: AGW. Humid bogs and springs, usually 

above 3500 m. Map 11. 

85. ARGENTINA NORTHWEST: Mendoza, between Portezuelo de Rahuí and valley of Chacaycó, F.Kurtz 

6071 (CORD); Arroyo del Papel, 3000 m, F.Kurtz 11127 (CORD); Mendoza, San Carlos, Laguna del 

Diamante, A.Ruíz Leal 14561 (K).


4. PATOSIA 

Patosia Buchenau, Bot. Jahrb. Syst. 12: 63 (1890) 

Distichia sect. Patosia (Buchenau) Kuntze in T.E. von Post & C.E.O.Kuntze, Lex. Gen. Phan. 182 (1903). 

T: Patosia clandestina (Phil.) Buchenau 

Perennial herbs, cushion-forming, glabrous. Stems ±regularly forked, dying back from the 

base, densely covered with persisting wilted leaves. Leaves regularly, densely, and 

alternately inserted along stem. Blade equal to or slightly shorter than sheath, flat or 

canaliculate; margins usually finely dentate. Inflorescence reduced to a single lateral subterminal 

flower. Plants dioecious or rarely with a few perfect flowers. Flower bracteole 

single. Tepals linear to lanceolate. Staminate flowers long pedicellate, exserted above the 

cushion; stamens 6, with long linear mucronate anthers on short filaments. Pistillate flowers 

sessile in axil of subapical leaf; ovary sessile; style long, filiform, with 3 filiform papillose 

stigmas. Capsule triseptate, sessile, narrowly ellipsoid. Seeds many; outer seed-coat spongy, 

irregularly wrinkled; inner seed-coat castaneous, hard. 

A single species distributed along the Andes from Bolivia to Chile and Argentina. 

1. Patosia clandestina (Phil.) Buchenau, Bot. Jahrb. Syst. 12 : 64 (1890), nom. conserv. 

propos. 

Rostkovia clandestina Phil., Linnaea 29: 76 (1857); Distichia clandestina (Phil.) Buchenau, Junc. S.-Amer. 6: 

370 (1879); Oxychloë clandestina (Phil.) Hauman, Anales Mus. Nac. Hist. Nat. Buenos Aires 27: 294 (1915). 

T: Chile, Coquimbo, Los Patos, R.A.Philippi s.n.; syn: SGO, n.v.; iso: K. 

Rostkovia brevifolia Phil., Linnaea 29: 76 (1857); Oxychloë brevifolia (Phil.) Buchenau, in A.H.R.Grisebach, 

Symb. Fl. Argent. 318 (1879). T: Chile, Laguna de Malvarco, Jan 1856, J.N.E.Germain s.n.; lecto: 

SGO63044, fide H.Balslev, Fl. Neotrop. Monogr. 68: 46 (1996); isolecto: SGO46499, n.v.; K; Cordillera de 

Sant Jago, R.A.Philippi s.n.; para: BREM, GOET, SGO46497. 

Patosia tucumanensis Castillon, Bol. Mus. Hist. Nat. Tucuman 7: 8 (1926); Patosia clandestina (Phil.) 

Buchenau var. tucumanensis (Castillon) Barros, Darwiniana 10: 295 (1953). T: Argentina, Tucumán, Valle 

de Tafí, Cerro Muños de la Piedra Pintada, Laguna de los Patos, 4100 m, 30 Dec 1916, L.Castillón 6039; holo: 

LIL, photo NY. 


Darwiniana 10: 290, fig. 2, 293, fig. 3 (1953); H.Balslev, Fl. Neotrop. Monogr. 68: 47, fig. 23 (1996). 

Perennials forming dense, very hard cushions up to several meters in diam. and 0.5 m high. 

Individual plants 5–15 cm long, ±regularly forked; stems covered with short spirally arranged 

wilted leaves and a few green leaves distally. Leaves 1.5–2 cm long, inserted 1–2 mm apart; 

sheaths as long as or longer than the blade, to 5 mm wide; auricles obtuse, sometimes fused 

and liguliform; blade linear, 0.5–0.7 mm wide, acuminate, proximally canaliculate, terete in 

the middle, distally flat with finely serrate margin; leaf seemingly jointed. Staminate flowers 

exserted on filiform pedicels 1–2 cm long; bracteole 4.0–4.5 mm long; tepals subequal, 

5–7 mm long; anthers 2.5–3.5 mm long, mucronate; filaments c. 0.2–0.3 mm long. Pistillate 

flowers sessile and hidden in axil with only stigmas exserted above the cushion; tepals linear 

to lanceolate, to 18 mm long; ovary ellipsoid, to 2 mm long, tapering to the filiform 10 mm 

long style; stigmas 3, filiform, to 8 mm long, papillose, not tapering, exserted. Capsule 

narrowly ellipsoid, acuminate, 10 × 2 mm. Seeds of irregular shape. Fig. 4. 

Confined to the high Andes from Bolivia to N and C Chile and from Jujuy to Neuquén, NW 

and SW Argentina. 83: BOL. 85: AGW, AGS, CLN, CLC. Alpine bogs, along streams, and 

in wet depressions, usually 3000–4000 m. Map 12. 

83. BOLIVIA: La Paz, Pacajes, Comanche, 4100 m, E.Asplund 2904 (UPS); H.Balslev & S.G.Beck 1085 

(AAU, LPB, NY); Cochabamba, Arque, road to Oruro, km 46, P.Rojas 856 (LPB). 85. ARGENTINA: 

Mendoza, San Carlos, Laguna del Diamante, 3000 m, A.Ruíz Leal 7240 (LIL); Neuquén, Copa-hué, 14 Feb 

1941, R.Pérez Moreau (BA37746, WU); Neuquén, Baño de Palau, 2250 m, Comber 297 (K). CHILE: 

Atacama, Copiapo, Sierra San Miguel, 3200 m, I.M.Johnston 4925 (F, GH, K, P, S, US); Cordillera de Maule, 

J.N.E.Germain (K). 

13

14 


Figure 4. Patosia clandestina. A, habit of clone (top) and a single shoot (right); B, leaf with 

basal sheath, channelled central part, and slightly dorsiventrally flattened distal part with 

finely serrate margin; C, staminate flower, note one bracteole; D, androecium, note 

mucronate anthers and rudimentary pistillode; E, single anther, note short filament and 

apical mucronate tip; F, pistillode from staminate flower, note three rudimentary stigmas; 

G, pistillate flower with protruding stigmas; H, gynoecium. (A–H, Johnston 4925). Scale bars: 

A = 5 cm; B–H = 1 mm. Drawn by H. Fukuda. Reproduced with permission, from H.Balslev, 

Fl. Neotrop. Monogr. 68: 47, fig. 23 (1996), © 1996, The New York Botanical Garden.


5. DISTICHIA 

Distichia Nees & Meyen, in F.J.F.Meyen, Observ. Bot. 128 (1843) 

Type: Distichia muscoides Nees & Meyen 

Goudotia Decne., Ann. Sci. Nat., Bot., Sér. 3, 4: 83 (1845); Distichia Nees & Meyen sect. Goudotia (Decne.) 

Benth. & Hook.f., Gen. Pl. 3: 367 (1883). T: Goudotia tolimensis Decne. 

Agapatea Steud., Bot. Zeitung (Berlin) 14: 391 (1856), nom.inval.; Agapatea Steud. ex Buchenau, Abh. 

Naturwiss. Vereine Bremen 4: 124 (1874), nom. inval. 

Perennials with branched suberect rhizomes, glabrous, cushion-forming. Stems ±regularly 

forked and covered with persisting leaves which are wilted in proximal part and live in the 

most distal part. Leaves regularly densely and strictly distichously inserted along stem, with 

a wide sheath clasping the leaf above; blade shorter than or equalling sheath, terete. 

Inflorescence reduced to a single lateral subterminal flower. Plants dioecious. Bracteoles 

2–4, minute, scarious. Tepals lanceolate, 4–10 mm long. Staminate flower long pedicellate. 

Stamens 6, with linear mucronate anthers about 10 times as long as the filaments. Pistillate 

flower short pedicellate; ovary on short gynophore; style with 3 filiform papillose stigmas. 

Capsule unilocular, lifted on a gynophore at ripening. Seeds many; outer seed-coat loose, 

irregularly wrinkled. 

Three species confined to the high Andes from Colombia and Ecuador to N Chile and 

N Argentina. 

1 Leaf apex bluntly acute, without hair-like extension 1. D. muscoides 

1: Leaf apex with hair-like extension or long acuminate 

2 Leaf blade gradually tapering to the tip; auricles absent 2. D. filamentosa 

2: Leaf blade linear, with tip long acuminate; auricles present 3. D. acicularis 

1. Distichia muscoides Nees & Meyen, in F.J.F.Meyen, Observ. Bot. 129 (1843) 

T: Peru, Puno, Pisacoma, 4500 m, Apr 1831, Meyen s.n.; holo: B, probably destroyed; syn: BR, P, photo QCA. 

Goudotia tolimensis Decne., Ann. Sci. Nat., Bot., Sér. 3, 4: 85 (1845); Distichia tolimensis (Decne.) Buchenau, 

Bot. Jahrb. Syst. 1: 141 (1880). T: Colombia, Tolima, 4600–5000 m, Feb 1828, J.Goudot s.n.; holo: P; iso: 

G, K, P. 

Gaimardia boliviana Pax, Repert. Spec. Nov. Regni Veg. 5: 225 (1908). T: Bolivia, La Paz, am Fusse des 

Murarata Swamps, 5000 m, Oct 1906, O.Buchtien 836; holo: B, probably destroyed; lecto: US, fide H.Balslev, 

Fl. Neotrop. Monogr. 68: 43 (1996); isolecto: UPS, photo NY. 

Agapatea peruviana Steud., in W.Lechler, Berberid. Amer. Austral. 56 (1857), nom.inval. Orig. coll.: Peru, 

Agapata, Jul 1854, W.Lechler 1954 (G, GOET, K, M, P, all p.p.). 

Illustrations: F.G.P.Buchenau, in H.G.A.Engler, Pflanzenr. (iv.36) 25: 34, fig. 27 (1906) as D. tolimensis; 

M.Barros, Darwiniana 10: 290, fig. 2 (1953); H.Balslev, Fl. Neotrop. Monogr. 68: 40, fig. 20A-C, 44, fig. 

22I-J (1996); H.Balslev, in [The Asahi Shinbun World of Plants; in Japanese] 122: 35 (1996). 

Perennials forming dense hard cushions up to several meters in diam. Single plants 5–15 cm 

long, with ±regularly forked shoots, covered with shor, strictly distichous persistent 

stramineous or rarely dark brown wilted leaves, with only 2 or 3 most distal leaves green and 

projecting above the cushion. Leaves 1–2 cm long, inserted 1–2 mm apart; sheaths usually 

two thirds of entire leaf length, open, wide, V-shaped in T.S., with narrow and scarious 

margins, almost completely clasping the sheath of the leaf above; blade bluntly acute. 

Flowers in a subterminal leaf axil. Tepals subequal, 4.5–7.5 mm long. Staminate flowers on 

filiform pedicels 1–2 cm long. Stamens 2.0–3.5 mm long; anthers 1.8–3.0 mm long; 

filaments c. 0.2–0.3 mm long. Pistillate flowers hidden in the sheath of the supporting leaf, 

with only stigmas projecting above the shoot apex; style filiform, exceeding tepals; stigmas 

half as long as style. Gynophore elongating at fruit ripening, lifting the capsule slightly 

above the cushion. Capsule ellipsoid to ovoid, apically acute to slightly apiculate or obtuse, 

5–7 × 2.5–3.5 mm, irregularly dehiscing, yellow-brown, unilocular. Seeds oblong, 1.0–1.5 × 

0.5–1.0 mm, covered by the thick white outer seed-coat. Fig. 1, 5. 

15

16 


Figure 5. Distichia acicularis. A, habit, staminate plant; B, shoot apex with staminate 

flower; C, leaf; D, shoot apex with stigmas protruding from pistillate flower; E, staminate 

flower; F, stamen; G, pistillate flower; H, gynoecium. (A-H, Laegaard 53236, AAU). 

Distichia muscoides. I, J, leaves, note conical blade and lacking auricles. (I, J, Laegaard 

53111, AAU). Scale bars: A = 5 cm; B, C, D, I, J = 10 mm; E, F = 5 mm; G, H = 5 mm. 

Drawn by K. Tind. Originally published in Nordic J. Bot. 6: 152, fig. 1 (1986), reproduced 

with permission, from H.Balslev, Fl. Neotrop. Monogr. 68: 44, fig. 22 (1996), © Lunds 

Botaniska Förening, © 1996, The New York Botanical Garden.

JUNCACEAE (Distichia) 

Confined to the Andes from Colombia to northern Argentina. 83: BOL, CLM, ECU, PER. 

85: AGW. In high-altitude paramo and puna vegetation, in boggy depressions and wet slopes 

with springs, usually from 4000–4600 m. The dense cushions are cut out and kept near 

houses, where they are used as fuel for cooking and heating by highland Indians. Map 13. 

83. BOLIVIA: La Paz, Lago Zongo, S side of Huayna Potosí, 4700 m, H.Balslev & S.G.Beck 1145 (AAU, 

CAS, LPB, NY); Oruro, Carangas, Sajama, 4200–5000 m, E.Asplund 3984 (NY, S, UPS). COLOMBIA: 

Caldas, Nevado del Ruiz, Termales, 4200–4500 m, J.-H.Humbert et al. 27076 (COL, GH, NY, S, SI, U, W); 

Santander, Páramo de Santurbán near Vetas, 3950–4160 m, E.P.Killip & Smith 17576 (C, COL, F, GH, NY, S). 

ECUADOR: Azuay, Lago Luspa, 3800 m, Ramsay 121 et al. (K); Chimborazo, E slope of Mt Chimborazo, 

4200 m, E.Asplund 7807 (AAU, BR, CAS, K, NY, S, UPS). PERU: Arequipa, Pichu Pichu, 4100 m, 

D.Stafford 691 (BM, K); Pasco, Huayllay, 4230–4610 m, Boeke 1098 (AAU, NY). 85. ARGENTINA 

NORTHWEST: Salta, Los Andes, 15 km N of Abra del Gallo, 4400 m, A.Charpin & L.J.Novara AC20775 (K); 

Tucumán, Cerro de las Animas, 4300 m, L.Castillón 6038 (LIL). 

2. Distichia filamentosa Buchenau, Junc. S.-Amer. 369 (1879) 

T: Bolivia, La Paz, Larecaja, Sorata, Laguna de Juriquana prope Anilaya, 4500 m, ii–iv 1860, G.Mandon 1443 

p.p.; syn: G, GH, K, NY, P, S, UPS. 

Agapatea filamentosa Buchenau, Abh. Naturwiss. Vereine Bremen 4: 124 (1874), nom. inval.; Distichia 

filamentosa Griseb., Symb. Fl. Argent. 318 (1879), nom. inval. Original material: Bolivia, La Paz, Sorata, San 

Pedro, 4500 m, 1860, G.Mandon 1443 p.p.; GOET. 

Perennials forming small cushions; shoots regularly forked, covered with short strictly 

distichous persistent stramineous or rarely dark brown wilted leaves, with only distal leaves 

green. Leaves c. 8–15 mm long; sheaths usually 6–8 mm long, c. 3–4 mm wide, with margins 

membranous; auricles ±not developed; blade usually 6–12 mm long, subconical-cylindric; tip 

with a 2–4 mm long filamentous extension. Flowers in a subterminal leaf axil. Tepals 

subequal, 4.5–7.5 mm long. Staminate flowers on 1–2 cm long filiform pedicels. Stamens 

2.0–3.5 mm long; anthers 1.8–3.0 mm long; filaments c. 0.2–0.3 mm long. Pistillate flowers 

hidden in the sheath of the supporting leaf, with only stigmas projecting above the shoot 

apex; style filiform, exceeding the tepals; stigmas half as long as style. Gynophore 

elongating at fruit ripening, lifting the capsule slightly above the cushion. Capsule ellipsoid 

to ovoid, subobtuse, mucronulate, 8–9 mm long. Seeds c. 1.5 mm long. 

A species known from the high Andes of La Paz province, Bolivia, from Arequipa in Peru 

and disjunctly from the province of Tarapaca in N Chile. 83: BOL, PER. 85: CLN. Flat 

shores of small ponds in alpine bogs, usually at 4000–4800 m. Map 14. 

83. BOLIVIA: La Paz, Sorata, San Pedro, 4500 m, G.Mandon 1443 p.p. (GOET); La Paz, Lago Zongo, S side 

of Huayna Potosí, 4700 m, H.Balslev & S.G.Beck 1146 (AAU, CAS, LPB, NY); La Paz, Mt Sorata, 4400 m, 

A.W.Hill 487 (B, K); La Paz, Murillo 6.4 km NE of La Cumbre, 4250 m, J.C.Solomon & Uehling 12133 (AAU, 

MO); La Paz, from El Cumbre to Coroico, E of the pass, 4800–4920 m, 18 Jul 1995, S.Laegaard 17669 

(AAU). PERU: Arequipa, Chucara Crater, 16 Jan 1999, 4700 m, B.Ruthsatz & S.G.Beck 10126 (AAU). 

85. CHILE: Tarapacá, Parinacota, 4820 m, B.Ruthsatz 8541 (AAU). 

3. Distichia acicularis Balslev & Laegaard, Nord. J. Bot. 6: 151 (1986) 

T: Ecuador, Azuay, Páramo de Soldados, 3700 m, 24 Oct 1984, S.Laegaard 53236; holo: AAU; iso: GB, K, 

NY, QCA, QCNE. 

Illustrations: H.Balslev & S.Laegaard, Nord. J. Bot. 6: 152, fig. 1A-H (1986); H.Balslev, Fl. Neotrop. 

Monogr. 68: 44, fig. 22A-H (1996). 

Map: H.Balslev & S.Laegaard, Nord. J. Bot. 6: 153, fig. 2 (1986). 

Perennials forming loose cushions up to 50 cm in diam. Individual plants consisting of 

±regularly forked shoots 5–15 cm long; proximal parts covered with stramineous wilted 

leaves; distal part with 6–8 green leaves. Leaves 1–2 cm long, completely clasping one 

another by the sheaths; sheaths equalling blades or shorter, 1.0–2.5 mm wide, U-shaped in 

T.S., with margins scarious; auricles pointed or rounded, 1–3 mm long, papery; blade linear, 

terete, 0.5–1.0 mm wide; apex drawn out into a long pointed tip. Staminate flower pedicels 

1.5–3.0 cm long; tepals linear-lanceolate, 5–8 mm long; stamens 2–6 mm long; gynoecium 

rudimentary, c. 1.5 mm long. Pistillate flower on pedicel c. 1 mm long; tepals linear- 

17

18 


lanceolate 7–10 mm long; ovary ovoid, c. 1 × 0.5 mm long; style 4–6 mm long; stigmas 5–7 

mm long. Fruit and seed unknown. Fig. 5. 

Found in the Western Cordillera of the Ecuadorian Andes at 1–3° S latitude. 83: ECU. High 

altitude spring bog areas, in seeping water, along streams, 3200–4200 m. Map 15. 

83. ECUADOR: Azuay, Páramo las Cajas W of Sayausí and Cuenca, 4100 m, H.Balslev 1481 (AAU, NY, 

QCA); Azuay, Páramo de Soldados, 3700–3800 m, S.Laegaard 53240 (UPS); Bolívar, road between Guaranda 

and Riobamba at km 10, 3200 m, S.Laegaard 51275 (AAU, MO, QCA); Chimborazo, Azul–Osogachi road, 

4000–4200 m, S.Laegaard 71009 (AAU, QCA); Cotopaxi, road between Latacunga and Quevedo, 18 km W of 

Pujilí, 3700 m, H.Balslev 1060 (AAU, NY, QCA); Tungurahua, road between Ambato and Guaranda, 3800 m, 

S.Laegaard 51265 (AAU, QCA, QCNE). 

6. LUZULA 

Luzula DC. in J.B.A.P. de M. de Lamarck & A.P. de Candolle, Fl. Franç., 3rd edn, 3: 158 

(1805), nom. cons. 

Juncoides Ség., Pl. Veron. 3: 88 (1754), nom. rejic. T: Luzula campestris (L.) DC. 

Luzula sect. Gymnodes Griseb., Spic. Fl. Rumel. 2: 405 (1846); Gymnodes (Griseb.) Fourr., Ann. Soc. Linn. 

Lyon, sér. 2, 17: 172 (1869); Juncoides sect. Gymnodes (Griseb.) Kuntze in T.E. von Post & C.E.O.Kuntze, 

Lex. Gen. Phan. 303 (1903). T: not designated. 

Nemorinia Fourr., Ann. Soc. Linn. Lyon, sér. 2, 17: 172 (1869). T: Luzula forsteri (Sm.) DC. [see Luzula 

subg. Pterodes] 

Perennial (rarely annual) herbs, rhizomatous, seldom cushion-forming, with sparsely to 

densely ciliate leaves, rarely leaves abaxially hairy; trichomes simple, many-celled. Leaves 

scale-like on rhizomes, bractlike below the inflorescence, usually narrowly linear, rarely 

linear-lanceolate, flat to concave; sheaths closed (rarely slightly dissected); auricles absent. 

Bracts leaf-like to scarious. Inflorescence diverse, usually many-flowered, either cymose 

with flowers borne singly and pseudo-paniculate, or racemose and then borne singly or in 

few-flowered clusters in much branched panicles or decompound anthelate inflorescences, or 

in spike-like clusters, either pedunculate or congested in many-flowered dense heads. Each 

flower subtended by a small bract at pedicel base and 1–2 subentire to fimbriate bracteoles 

just below flower. Flowers hermaphrodite, perfect. Tepals 6, equal to ±unequal, ±lanceolate 

(linear-lanceolate to ovate-lanceolate), glumaceous, glabrous or ciliate-fimbriate. Stamens 3–6; 

anthers oblong to linear; filaments filiform to linear; ovary sessile; style thin, 0.1–c. 4 mm 

long; stigmas 3, suberect, twining. Capsule ±ovoid-trigonous, unilocular. Seeds 3, with or 

without appendages (caruncula); outer seed-coat tight, usually hyaline, less often whitish or 

brownish. x = 6 (sometimes changed by fragmentation, mostly simultaneous (agmatoploidy), 

or fusion (x = 3)), fide H.Nordenskiöld, Hereditas 37: 325–355 (1951), 42: 7–73 (1956). 

A genus of 115 species, almost cosmopolitan, much rarer in the tropics, with major diversity 

centres in SW Europe, the Far East, W North America, temperate South America (the Andes) 

and Australia and New Zealand. Not important economically. 

F.G.P.Buchenau, Monographia Juncacearum, Bot. Jahrb. Syst. 12: 1–495 (1890); 

F.G.P.Buchenau, Juncaceae, in H.G.A.Engler, Pflanzenr. (iv.36) Heft 25: 1–284 (1906); 

H.Nordenskiöld, Cyto-taxonomical studies in the genus Luzula I, II, Hereditas 37: 325–355 

(1951), 42: 7–73 (1956); L.Hämet-Ahti, A synopsis of the species of Luzula, subgenus 

Anthelaea Griseb. (Juncaceae) indigenous in North America, Ann. Bot. Fenn. 8: 368–381 

(1971); V.S.Novikov, Konspekt sistemy roda Luzula DC. (Juncaceae), Byull. Moskovsk. 

Obshch. Isp. Prir., Otd. Biol. 95(6): 63–70 (1990); J.Kirschner, An account of natural 

hybridization within Luzula sect. Luzula (Juncaceae) in Europe, Preslia 63: 81–112 (1991); 

J.Kirschner, Karyological differentiation of Luzula sect. Luzula in Europe, Thaiszia 2: 11–39 

(1992); J.Kirschner, Taxonomic survey of Luzula sect. Luzula (Juncaceae) in Europe, Folia 

Geobot. Phytotax. 28: 141–182 (1993). 

The genus is characterized by chromosomes with diffuse or polycentric centromeres, and, 

during its evolution, it undergoes (often repeatedly) a simultaneous chromosome

JUNCACEAE (Luzula) 

fragmentation. The phenomenon is called agmatoploidy; in consequence, a mere 

chromosome number does not give information about the ploidy level. Chromosome numbers 

are therefore given with data on karyotype; unfragmented chromosomes are designated AL, 

half-sized chromosomes BL and quarter-sized ones are called CL. 

For the compilation of descriptions, seeds were measured in dorsal view, seed length 

excludes the appendage, and lower flowers of clusters were examined for floral traits (tepals 

and capsule segments first liberated from inflorescence). 

KEY TO SUBGENERA AND SECTIONS 

1 Annuals; inflorescence cymose; n = 3 subg. 1. Marlenia 

1: Perennials; inflorescence racemose; n = 6 

2 Seed appendage apical (distal), large subg. 3. Pterodes 

2: Seed appendage basal or absent, but distal end of seed sometimes with a 

minute pale patch of loose outer seed-coat 

3 Flowers usually borne singly in the inflorescence, rarely some in 

groups of 2–5; inflorescence much branched sect. 4. Diprophyllatae 

3: Flowers in dense pedunculate clusters or wholly congested, or in 

groups of (2) 3–5 (to many) flowers in the whole inflorescence; 

inflorescence sparsely branched, unbranched or much branched 

4 Leaf tip obtuse to subobtuse, often slightly swollen, rarely with a 

spinuliform mucro 

5 Leaf tip obtuse, not swollen, with a spinuliform mucro 49. Luzula nivalis 

5: Leaf tip obtuse or subobtuse, slightly swollen or not swollen, 

without spinuliform mucro sect. 7. Luzula 

4: Leaf tip acute to acuminate, not swollen 

6 Inflorescence usually nodding, less often erect (then stamens 3 or 

tepals ciliate; high Andes), spike-like or paniculate (a panicle of 

±dense spike-like clusters), clusters congested or interrupted at base sect. 5. Alpinae 

6: Inflorescence erect to suberect, if slightly nodding then much 

branched, loose, of (2) 3–5 (20)-flowered groups, anthelate (or 

corymbose) or umbelloid, of distinct clusters (if congested then 

stamens 6, tepals glabrous; N hemisphere) 

7 Inflorescence much branched, (doubly) anthelate, terminal groups 

mostly of 2–4 (8) flowers, head-like sect. 1. Anthelaea 

7: Inflorescence sparsely branched or unbranched, umbelloid to 

subcongested, of distinct pedunculate clusters (or wholly 

contracted), clusters 3–8 (–many)-flowered 

8 Tepals ±membranous; seeds ±globose, smooth sect. 2. Atlanticae 

8: Tepals coloured (usually castaneous-brown); seeds broadly 

ovoid to ellipsoid, surface sculptured 

9 Seeds 1.7–1.9 × 1.3–1.4 mm; rhizome thick, nodulose sect. 3. Nodulosae 

9: Seeds 0.9–1.3 × 0.5–0.3 mm; rhizome thin or indistinct 

10 Leaf margin smooth 91. L. acutifolia 

10: Leaf margin densely papillose-serrulate sect. 6. Thyrsanochlamydeae 

19

20 


Subg. 1. MARLENIA 

Luzula subg. Marlenia Ebinger, Brittonia 15: 173 (1963) 

Ebingeria Chrtek & Křísa, Preslia 46: 210 (1974). T: Luzula elegans Lowe 

Luzula sect. Triprophyllatae Satake in T.Nakai & M.Honda, Nov. Fl. Jap. 1: 25 (1938). T: Luzula purpurea 

E.Mey. 

Annuals. Leaves acute; apex extending to a hair, sometimes two hairs; leaf margin without 

sclerenchyma. Inflorescence cymose; flowers borne singly, terminal. Endodermis with walls 

evenly thickened (O-shaped in T.S.). Seeds smooth; appendage absent. n = 3. 

A monotypic, remarkably distinct subgenus. The only species, Luzula elegans, is native to 

Macaronesia and introduced to Portugal. 

1. Luzula elegans Lowe, Trans. Cambridge Philos. Soc. 6: 532 (1838) 

Ebingeria elegans (Lowe) Chrtek & Křísa, Preslia 46: 211 (1974). T: [Madeira] ‘in rupibus convallium 

Maderae murisque rarior’; [no locality] 8 May 1838, R.T.Lowe 646; syn: BM. 

Luzula purpurea Link [in C.L. von Buch] ex E.Mey., Linnaea 22: 397 (1849); Juncoides purpurea (E.Mey.) 

Kuntze, Revis. Gen. Pl. 2: 725 (1891). T: [Canary Is.] La Palma, Orotava, C.L.Buch; syn: n.v.; Tenerife, 

Forsyth; syn: K, n.v.; Tenerife, Lenormand; syn: LE, n.v. 

Luzula berthelotii Nees ex Kunth, Enum. Pl. 3: 298 (1841). T: ‘In sylva de las Mercedes Insulae 

Teneriffae’; Tenerife, P.B.Webb; holo: B, destroyed; iso: K, BM. 

Illustration: F.G.P.Buchenau, in H.G.A.Engler, Pflanzenr. (iv.36) 25: 50, fig. 40 (1906); J.E.Ebinger, 

Brittonia 15: 170, fig. 1, 3, 172, fig. 5, 8, 9 (1963). 

Map: H.Meusel, E.Jäger & E.Weinert, Vergl. Chorol. Zentraleur. Fl. 1: 87 (1965). 

Annuals, usually 1–2-stemmed, 8–17 (–25) cm. Basal leaves several, to 12 cm long, 1.2–2.8 mm 

wide. Cauline leaves 2–4, to 10 cm long, to c. 2.5–3.0 mm wide; margins ±smooth, sparsely 

hairy, with hairs long, especially near sheath mouth; apex acute, usually ±extended to a 

single or sometimes two hairs. Inflorescence bract herbaceous, c. 2–4 cm long, surrounding 

pseudorhachis; middle bracts purplish-membranous with scarious ciliate margins. Bracteoles 

purplish-membranous, ovate, acuminate, c. 2.0–2.5 mm long. Inflorescence a loose divaricate 

panicle usually 5–13 cm long, almost evenly branched or distally denser; branches thin, smooth; 

flowers borne singly. Tepals ±equal or outer slightly longer, ±ovate, with outer tepals purplish 

with decreasing colour intensity towards membranous margins, (2.4–) 2.7–2.9 (–3.2) mm long, 

acute and mucronate; midrib usually distinctly papillose near apex; inner tepals wholly 

translucent-membranous or midrib purplish; apex usually emarginate and mucronate. 

Stamens 6; anthers 0.3–0.5 mm long; filaments 0.4–0.8 mm long; style 0.2–0.3 mm long; 

stigmas 0.3–0.5 mm long, not exserted. Capsule ovoid-subglobose, pale brown to pale 

purplish, thin-walled, ±acute to subobtuse; segments 1.4–1.8 mm long, c. 1.3–1.5 mm wide; 

mucro short. Seeds subglobular 0.8–0.9 × 0.6–0.7 mm long, smooth, shining, pale 

castaneous-brown; appendage absent. 2n=6, fide N.Malheiros & D.Castro, Nature 160: 156 

(1947); H.Nordenskiöld, Hereditas 37: 327 (1951). 

Native to Madeira, Canary Is. (Gran Canaria, Tenerife, Gomera, Hierro, La Palma), 

naturalized in Portugal (Estremadura and Coimbra). 12: por. 21: CNY, MDR. On rocks, walls, 

dry slopes, along paths. Map 16. 

12. PORTUGAL: Coimbra, 1879, Moller (WU); Estremadura, Sacavém, Quinta dos Almastéres, 26 Apr 1946, 

Fontés & Rainha (LD). 21. CANARY IS: Tenerife, El Boquerón, 6 Mar 1988, R.Lindberg (S); Tenerife, 

Güímar, Barranco del Río, 25 Apr 1933, E.Asplund (S, UPS); Gran Canaria, between San Mateo and 

Valsequillo, 1250 m, C.C.Townsend 78/2 (K); Hierro, from Valverde to Frontera, C.E.Jarvis & Murphy 298 

(BM); Hierro, El Golfo, Risco de Jinama, J.F.N.Bornmüller [Pl. Exs. Canar.] 2907 (LD, P, PR); 

La Palma, Barranco de las Angustias, 1000 m, Mar 1906, C.-J.M.Pitard 694 (SAM). MADEIRA: Madeira, 

between Ribeira Frio and Balcoes, 1100 m, 29 Apr 1952, G.Een & Persson (S); Madeira, São Roque, 

G.Mandon [Pl. Mader.] 248 (K, P, PR). 

In many respects close to the N African Luzula atlantica (2n=12). The chromosome number 

of L. elegans is unique in the genus, and there is a plausible suggestion that the number is a 

result of chromosome fusion.


Subg. 2. LUZULA 

Luzula B. Stachyophora C.Koch, Linnaea 21: 625 (1848). T: not designated [sections Alpinae, Anthelaea 

and Luzula represented]. 

Perennials. Leaves with acute, mucronate or obtuse tips. Inflorescence racemose; flowers 

borne singly in much branched decompound inflorescences, or in few- to many-flowered 

head-like groups, or in pedunculate clusters, sometimes wholly congested. Endodermis of 

cells with U-shaped thickened walls in T.S. Seeds finely reticulate or smooth; appendages 

basal or ±absent. 

Sect. 1. ANTHELAEA 

Luzula sect. Anthelaea Griseb., Spic. Fl. Rumel. 2: 404 (1846) 

Luzula subg. Anthelaea (Griseb.) Buchenau, Bot. Jahrb. Syst. 1: 113 (1880); Juncoides sect. Anthelaea 

(Griseb.) Kuntze, in T.E. von Post & C.E.O.Kuntze, Lex. Gen. Phan. 303 (1903); Luzula subsect. Anthelaea 

[Griseb.] Novikov, Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 95(6): 66 (1990). T: Luzula maxima 

(Reichard) DC.; lecto, designated here. 

Luzula maxima (Reichard) DC., Preslia 46: 211 (1974). T: Luzula luzuloides (Lam.) Dandy & E.Willm. 

Perennials, usually with ascending branched rhizomes. Leaves with acute tips. Lower bract 

herbaceous. Inflorescence decompound, an anthela of anthelas. Terminal clusters usually of 

2–4 flowers, tepals usually 2.5–5.5 mm long. Seed appendage basal, short, 0.1–0.2 mm, 

fibrillate. 

A mostly European section with 10 species centred to the Alps, SW Europe and Macaronesia. 

Most of the species are endemics confined to small areas. 

1 Tepals conspicuously unequal, with inner ones longer 

2 Tepals white 

3 Tepals (2.0–) 2.3–3.2 mm long 9. L. luzuloides 

3: Tepals 4.2–5.5 mm long 6. L. nivea 

2: Tepals yellow, purplish, coppery-coloured or grey-brownish 

4 Tepals yellow; flower clusters usually 10–15-flowered, 

glomeruliform; tepals up to 3.2 mm long 8. L. lutea 

4: Tepals purplish, brownish-purplish or coppery-coloured (later 

sometimes becoming grey-brownish); flower clusters 2–8-flowered; 

tepals usually longer than 3.5 mm 

5 Tepals 4.4–5.0 mm, purplish or bright copper-coloured; style 

1.4–1.6 mm long; stigmas 1.0–1.4 mm long 3. L. purpureosplendens 

5: Tepals 3.0–4.2 mm, copper-coloured or brownish-purplish; style 

0.8–1.2 mm long; stigmas c. 0.6–1.1 mm long 9. L. luzuloides 

1: Tepals equal or subequal 

6 Bracteoles linear-lanceolate, at least 3.5 mm long; tepals 5.0–6.5 mm 

long. [Madeira] 2. L. seubertii 

6: Bracteoles ovate or ovate-lanceolate, up to 3 mm long; tepals up to 

5 mm long, usually much shorter 

7 Flowers white, dirty white, cream-white or whitish membranous 

8 Inflorescence dense, ±subcontracted; flower clusters usually 

(3–) 5–15-flowered 

9 Basal leaves 3–4 mm wide; capsules 1.9–2.2 mm long including 

mucro; abaxial leaf surface hairy or glabrous, with margins ciliate 4. L. lactea 

21

22 


9: Basal leaves 13–19 mm wide; capsules c. 2.5 mm including mucro; 

abaxial leaf surface glabrous, with margins often subglabrous 5. L. canariensis 

8: Inflorescence lax or ±dense, ±evenly branched, not subcontracted; 

flower clusters 2–5 (–7)-flowered 

10 Anthers bifurcate in distal 1/5– 1/3; bracteoles 2.0–2.5 mm long 7. L. pedemontana 

10: Anthers not bifurcate; bracteoles 1.0–1.5 mm long 9. L. luzuloides 

7: Flowers ±brown, castaneous-brown, brownish copper-coloured or 

membranous with castaneous-brown midrib 

11 Style 0.8–1.2 mm long; stigmas less than 1.5 mm long; tepals with 

purplish or coppery colour/tinge 

11: Style 1.6–2.2 mm long; stigmas longer than 2 mm; tepals without 

9. L. luzuloides 

purplish or coppery colour/tinge 10. L. sylvatica 

2. Luzula seubertii Lowe, J. Bot. (Hooker) 8: 300 (1856) 

Juncoides seubertii (Lowe) Kuntze, Revis. Gen. Pl. 2: 724 (1891). T: ‘In salebrosis rupestribus umbrosis 

Maderae’. Madeira, Serra do Seixal, Cruzinhas, 23 Jul 1855, R.T.Lowe; lecto: K, designated here; isolecto: P; 

other authentic specimens from type locality: K, LD. 

Illustration: M.J.Cannon, in J.R.Press & M.J.Short, Fl. Madeira, plate 53, fig. 4 (1994). 


Perennials, usually 40–80 cm tall, densely ciliate, loosely caespitose; rhizome creeping to 

ascending, long, often with dark fibrous cover; stolons ascending. Basal leaves few, up to 

35–40 cm long, usually (6–) 8–11 mm wide; sheaths brown. Cauline leaves 3–5; upper leaves 

usually 5–8 cm long, 3–5 mm wide. All leaves with acuminate tip; margins ±smooth. Lower 

bracts ±herbaceous, 2–7 cm long; apex long acuminate, much shorter than inflorescence. 

Inflorescence decompound lax, with slightly drooping rich anthela to 15 cm long; branches 

smooth; terminal parts of inflorescence branches anthelate; terminal flower groups loose 

(1–4-flowered); peduncle visible and usually (0.5–) 1.5–5.0 (–8.0) mm long. Bracteoles 

linear-lanceolate, long acuminate, carinate, membranous to pale straw-brown, sparsely long 

ciliate, 3.5–5.5 mm long. Tepals linear-lanceolate, gradually tapering to long acuminate tip, 

equal to subequal, usually (5.0–) 5.5–6.5 mm long, pale straw-brown with membranous 

margins. Stamens 6; anthers 1.3–1.6 mm long; filaments c. 1.0 mm long; ovary 1.2–1.3 mm 

long; style 1.0–1.3 mm long; stigmas 1.5–2.8 mm long. Capsule pale brown, ±shining, ovoid, 

±acuminate; mucro c. 0.3–0.5 mm long; capsule segments 2.8–3.0 × 1.6–1.9 mm. Seeds 

oblong-ovoid, dark brown, 1.3–1.4 mm long, 0.9–1.0 mm wide; appendage brownish, 

0.1–0.2 mm long, fibrillate. 2n=12, fide N.Malheiros & A.Gardé, Agron. Lusit. 9: 76 (1947). 

Endemic to mountains of Madeira. 21: MDR. Dry mountain grassland and shrubby slopes, 

usually on rocky ground. Map 17. 

21. MADEIRA: Serra do Seixal, Cruzinhas, 18 Jul 1850, R.T.Lowe (LD, K); Serra do Seixal, Cruzinhas, 

30 May 1863, R.T.Lowe (BM, K); Mt Torrinhos, 29 Jun 1828, P.B.Webb (K); Boco de Boroaca, Pico dos 

Arrieros, 1200–1500 m, G.Mandon [Pl. Mader.] 247 (BM, K, P, PR, W); Levada do M. Medonho, Aug 1892, 

Murray (BM). 

3. Luzula purpureosplendens Seub., Fl. Azor. 23 (1844) 

Luzula elegans Guthnick [in C.F.F.Hochstetter] ex E.Mey., Linnaea 22: 392 (1849), nom. illeg.; Luzula 

purpurea H.C.Watson, London J. Bot. 3: 608 (1844), nom. illeg. T: Azores, H.J.Guthnick [in 

C.F.Hochstetter, Herb. Azor.] 126; lecto: K, designated here; isolecto: BM, P, W. 

Luzula azorica H.C.Watson, London J. Bot. 2: 408 (1843), nom. inval. Authentic material: Azores, 1842, 

H.C.Watson; K. 

Illustration: M.A.Seubert, Fl. Azor., Plate iv, fig. 2. (1844). 

Perennials, 25–45 (–60) cm tall, caespitose to loosely caespitose; rhizome ascending, 

branched; stolons numerous, ascending, with bases covered with brown to purplish 

castaneous-brown cataphylls. Basal leaves numerous, usually 15–22 cm long, 3–4.5 (–6) mm


wide. Cauline leaves 1.5–3.5 (–8.0) cm long. All leaves subulate; margins ±smooth, ciliate to 

glabrescent, slightly swollen. Lower bract subherbaceous, erect, 1.2–2.5 cm long, much 

shorter than inflorescence. Inflorescence decompound, ±erect, usually 4–5 × 3–4 cm, dense, 

with each main branch bearing anthela of short pedunculate usually 2–6-flowered clusters. 

Bracteoles purplish membranous to pale grey, ±lanceolate, sparsely ciliate, 2.5–3.0 mm long. 

Tepals narrowly oblong-lanceolate, glabrous, ±unequal, with inner tepals often conspicuously 

longer, 4.4–5.0 mm long, purplish-coppery with hyaline margin, later becoming greyish pale 

brown; midrib often pale. Stamens 6; anthers 1.6–1.9 mm long; filaments c. 0.5 mm long; 

style c. 1.4–1.6 mm long; stigmas 1.0–1.4 mm long. Capsule subglobular, abruptly narrowing 

to mucro 0.3–0.5 mm long, dark castaneous-brown, shining; capsule segments 1.8–2.0 × 

c. 1.2 mm including mucro. Seeds dark brown, oblong-ovoid, c. 0.9 mm long, 0.7–0.8 mm 

wide; appendage indistinct, c. 0.1 mm long, fibrillate. 

Endemic to the Azores. 21: AZO. Open broad-leaved woodlands, shrubby slopes, deforested 

mountain summit areas. Map 18. 

21. AZORES: São Miguel, Hunt (K, UPS, W); São Miguel, Lagoa das Sete Citades, O.Degener & I.Degener 

36253 (S); São Miguel, Ribeira Grande, Lagoa do Fogo, 755 m, 14 May 2000, J.Kirschner (PRA); 

São Miguel, Furnas, Terra Nostra Park, 14 May 2000, J.Kirschner (PRA); Pico, between Lages and Cais do 

Pico, 11 May 1937, H.Persson (S); Pico, Santa Luzia, 400 m, M.L.Gonçalves 1105 (BM); ‘in sylvaticis insul. 

Azoricarum’, H.J.Guthnick s.n. (W); Flores, Rocha do Junco, M.L.Gonçalves 4264 (BM); Faial, Caldeira 

Grande, 4 May 1937, H.Persson s.n. (H). 

4. Luzula lactea (Link) E.Mey., Syn. Luzul. 15 (1823) 

Juncus lacteus Link, J. Bot. (Schrader) 2: 316 (1799); Juncus brevifolius Link ex Rostk., De Junco 35 (1801); 

Luzula brevifolia (Rostk.) Poir., Encycl., Suppl. 3: 532 (1813); Juncoides lactea (Link) Kuntze, Revis. Gen. 

Pl. 2: 724 (1891). T: [Portugal] ‘zwischen den Felsen am Gerez’ [Serra do Gerês], J.H.F.Link; holo: B-W. 

Maps: P.Montserrat, Anales Inst. Bot. Cavanilles 21(2): map 2 (1964); H.Meusel, E.Jäger & E.Weinert, Vergl. 

Chorol. Zentraleur. Fl. 1: 87 (1965). 

Perennials, usually 30–55 cm tall, loosely caespitose; rhizome ±creeping; stolons subterranean, 

±ascending, to 12 cm long, with brown cataphylls and sparse fibrous cover. Basal leaves 

usually 8–15 cm long, often circinate, flat, c. 3–4 mm wide; cauline leaves 3–5, usually 3–7 cm 

long, c. 2 mm wide; sheaths dissected, ±dilated above, membranous bordered, ±abruptly 

narrowing to blade. All leaves with subulate apex; margins papillose-serrulate. Lower bract 

usually 2.0–3.5 cm long, subherbaceous, shorter or rarely ±longer than inflorescence. 

Inflorescence decompound, dense, usually 2–5 × 1.5–5 cm, with each main branch bearing 

dense anthela of 5–30 glomerate (usually 5–15-flowered) clusters. Bracteoles ±membranous, 

transparent, ovate-lanceolate, 2.5–3.0 mm long, sparsely ciliate. Tepals ±equal, glabrous, 

oblong-lanceolate, (4.0–) 4.2–4.6 (–5.2) mm long, ±white, subtransparent-ivory white to dirty 

whitish; midrib sometimes prominent. Stamens 6; anthers 1.1–1.5 (–1.8) mm long; filaments 

1.5–1.9 mm long; style 1.9–2.2 mm long; stigmas 0.6–1.1 mm long. Capsule narrowly ovoid, 

acuminate, mucronate, castaneous-brown, shining; capsule segments 1.9–2.2 × c. 1.0 mm, 

including mucro 0.2–0.4 mm long. Seeds ellipsoidal, pale brown, shining, ±paler apically, 

1.0–1.1 mm long, c. 0.6 mm wide; appendage 0.1–0.2 mm long, fibrillate. 

Confined to the mountains of Spain and Portugal. Two taxa are recognized on the basis of 

the leaf indumentum; intermediates in the southern part of the geographical range, and 

occasional coexistence of the two varieties within populations, suggest that this character is 

of doubtful taxonomic value. A late flowering species. Reported from Azores but no material 

seen (probably a misidentification of a late, ±greyish-flowered L. purpureosplendens). 

Abaxial surface of leaves glabrous, with margins ciliate 4a. var. lactea 

Abaxial surface of (at least some) leaves densely hairy 4b. var. velutina 

23

24 


4a. Luzula lactea (Link) E.Mey. var. lactea 

Juncus stoechadanthos Brot., Fl. Lusit. 1: 516 (1804). T: Portugal, ‘in summis montosis Gerez’, J.H.F.Link; 

holo: B, destroyed, probably a duplicate of the type of L. lactea. 

Illustration: F.G.P.Buchenau, in H.G.A.Engler, Pflanzenr. (iv.36) 25: 57, fig. 43 (1906). 

Leaf margin ciliate to glabrescent; abaxial surface glabrous. Basal leaves arcuate to straight, 

less often circinate. 2n=12, fide N.Malheiros & A.Gardé, Agron. Lusit. 9: 76 (1947). 

Mountains of NW and central Spain and northernmost Portugal. 12: POR, SPA. Open dry 

oak woodlands and clearings. Map 19. 

12. PORTUGAL: Serra da Estrela, Facarão, 1500 m, M.Ferreira [A.Kneucker, Cyper. Junc. Exs.] 210 (PR, W, 

WU). SPAIN: Soria, Sierra de Urbión, 5 Jul 1958, N.Y.Sandwith 5366 (K); Puerto de Leitariegos, 7 Aug 

1835, M.C.Durieu [Pl Select. Hisp. Lusit.] 214 (K, W); Sierra de Guadarrama, Cercedilla, C.Vicioso (LD, M); 

Sierra de Guadarrama, Cercedilla, 18 Jul 1924, Ginzberger (WU); Sierra de Guadarrama, Puerto de 

Navacerrada, 1800 m, K.H.Rechinger 62610 (W); Sierra de Guadarrama, Puerto de Navacerrada, E.Levier 

(W 13429); León, Cordillera Cantabrica, between Valverde and Puerto de Vegarada, Cerulleda, 1200 m, 

B. de Retz 76163 (LD). 

Intermediates between the two varieties also occur in Serra do Gerez (LD). The type material 

of L. lactea in B-W has only been seen as a photograph. 

4b. Luzula lactea var. velutina (Lange) Cout., Bol. Soc. Brot. 8: 122 (1890) 

Luzula velutina Lange, Vidensk. Meddel. Dansk. Naturlist. Foren. Kjöbenhavn 1881: 93 (1881); Luzula lactea 

subsp. velutina (Lange) Nyman, Consp. Fl. Eur. 4: 751 (1882). T: ‘In monte Serra do Ratico c. Murca 

Lusitaniae’ [Ponte do Ratinho, Murça], Jul. 1880, M.Ferreira; holo: C, n.v. 

Leaf margin usually ciliate; abaxial surface usually densely hairy. Basal leaves usually 

distinctly circinate. 

Mountains of west-central Spain and east-central Portugal. 12: POR, SPA. Open dry oak 

woodlands and clearings. Map 20. 

12. PORTUGAL: Beira Baixa, Serra da Estrella, Facarão, 1500 m, M.Ferreira [A.Kneucker, Cyper. Junc. 

Exs.] 178 (PR, PRA, W); Serra da Estrella, Poio Negro, Pimentel [Fl. Lusit. Exs.] 1838 (LD, PR); Serra da 

Estrella, Penha Douradas, 1550 m, 13–25 Jun 1987, A.Polatschek (W); Beira Baixa, Serra de Gardunha, Valle 

de Prazeres, 900 m, 16 Jun 1938, W.Rothmaler (S); Ponte do Ratinho, between Murça and Palheiros, 

A.Fernandes et al. 5423 (UPS); Guarda, Videmonte, 16 Jun 1953, A.Fernandes et al. 4386 (LD, UPS); Alto 

Alentejo, Portalegre, 1000 m, J.V.C.Malato-Beliz et al. 16310 (LD). 

5. Luzula canariensis Poir., Encycl., Suppl. 3: 532 (1813) 

Juncoides canariensis (Poir.) Kuntze, Revis. Gen. Pl. 2: 724 (1891). T: Canary Is., Tenerife, Laguna, 

Broussonnet; holo: P-DESF, n.v. 


Perennials 35–90 cm tall, loosely caespitose, subglabrous; rhizome long, creeping to 

ascending; stolons present; plant base often purplish. Stem thick, c. 2–3 mm in diam. Basal 

leaves usually 18–30 cm long, 13–19 mm wide, evenly narrowing; cauline leaves usually 

4–5, up to 18 cm long and 13 mm wide. All leaves acute; margins papillose-serrulate; 

sheaths ±dissected, with mouth ciliate. Lower bract 2–3 cm long, herbaceous, shorter than 

inflorescence. Inflorescence decompound, ±dense, usually 1.5–8 × 2.5 × 6 cm; main branches 

with anthelas of 3–8-flowered clusters. Bracteoles ovate-lanceolate, 2.0–2.5 mm long, 

membranous-dirty white, sparsely ciliate. Tepals ±equal, glabrous, oblong-lanceolate, usually 

(3.0–) 3.5–4.2 mm long, dirty white (whitish-membranous with pale straw-brown base and 

1–3 prominent veins). Stamens 6; anthers 1.1–1.3 mm long; filaments c. 1.0–1.1 mm long; 

style c. 1.0 mm long; stigmas 0.9–1.3 mm long. Capsule ±ovoid, acuminate, mucronate, pale 

brown; capsule segments c. 2.5 × 1.5–1.6 mm long, including mucro c. 0.5 mm long. Seeds 

ellipsoidal, 1.1–1.3 mm long, 0.6–0.7 mm wide; appendage c. 0.1 mm long, probably not or 

indistinctly fibrillate. 2n=12, fide L.Borgen, Check-List Chromos. Macarones. Vasc. Pl. 30 

(1977).


Restricted to Canary Is., islands of Gomera and Tenerife; formerly also reported from Gran 

Canaria but material not seen. 21: CNY. Found in broad-leaved woodlands. Map 21. 

21. CANARY IS.: Tenerife, Las Mercedes, E.Bourgeau [Pl. Canar.] 1553 (K, PR, UPS); Tenerife, Las 

Mercedes, R.T.Lowe 245 (S); Tenerife, Las Mercedes, J.F.N.Bornmüller [Pl. Exs. Canar.] 1273 (LD, PR); 

Tenerife, Las Mercedes, F.Ehrendorfer et al. 096 (W); Tenerife, Las Mercedes, between Puerto del Bailadero 

and Los Inglés, 900 m, B. de Retz 81442 (LD); Tenerife, Anaga, Las Hiedras, 1 Jun 1972, Beltrán (K, TFC); 

Mt Anaga, O.Burchard [Pl. Canar. Exs.] 158 (S); Tenerife, Taganana, E.H.Jörgensen 2968 (UPS); Gomera, 

Monte del Cedro, top of Barranco del Hermigua, 900 m, C.E.Jarvis 602 (BM); Monte del Cedro, Garajonay, 

1300 m, E.Wängsjö & G.Wängsjö 4923 (LD) 

6. Luzula nivea (Nathh.) DC., in J.B.A.P. de M. de Lamarck & A.P. de Candolle, Fl. 

Franç., 3rd edn, 3: 158 (1805) 

Juncus niveus Nathh., in C.Linnaeus, Amoen. Acad. 4: 481 (1756); Juncoides nivea (Nathh.) Kuntze, Revis. 

Gen. Pl. 2: 724 (1891). T: Magnol, Bot. Monspel., no 319. 

Luzula nivea var. livida Desv., J. Bot. (Desvaux) 1: 151 (1809); Luzula nivea subvar. livida (Desv.) Rouy, Fl. 

France 13: 260 (1912). T: not designated. 

Luzula nivea var. rubella Mert. & W.D.J.Koch, Deutschl. Fl. (Röhling) 2: 600 (1826); Luzula nivea var. 

rubescens Favrat, in A.Gremli, Excursionsfl. Schweiz, 5th edn, 419 (1885); Luzula nivea subvar. rubella 

(Mert. & W.D.J.Koch) Rouy, Fl. France 13: 261 (1912). T: Austria, ‘Villacher Alpe’, L.C. von Vest; holo: n.v. 

Luzula nivea var. minuta Beyer, Verh. Bot. Vereins. Prov. Brandenburg 41: xvii (1900). T: Switzerland, 

Cadenazzo, Mte. Ceneri, 31 Jul 1883, R.Beyer; syn: W; Italy, Piemont, Val Pesio, Certosa, R.Beyer; syn: n.v.; 

Switzerland, Mt Generoso, Scherer; syn: B, destroyed; Switzerland, Saléve, Guillemin; syn: B, destroyed. 

Luzula nivea b. [unranked] angustata Asch. & Graebn., Syn. Mitteleur. Fl. 2(2): 507 (1904); Luzula nivea var. 

angustata (Asch. & Graebn.) Rouy, Fl. France 13: 261 (1912). T: not designated. 

Illustration: H.E.Hess, E.Landolt & R.Hirzel, Fl. Schweiz 1: 517 (1967). 


Perennials usually 45–85 cm tall, loosely caespitose, ciliate; rhizome ascending, often with 

fibrous cover; stolons ascending, with base covered with brownish cataphylls. Basal leaves 

usually (15–) 20–30 cm long, usually 3–4 (–5) mm wide; cauline leaves 3–5, to 30 cm long, 

often exceeding inflorescence. All leaves acute, papillose-serrate (basal leaves often ±smooth). 

Lower bract herbaceous, usually up to 10–12 cm long, longer than inflorescence. Inflorescence 

decompound, typically 3–5 × 3–5 cm; main branches short, each with a ±dense anthelate 

5–12 (–20)-flowered cluster. Bracteoles ovate to ovate-lanceolate, whitish-scarious, c. 1.5 mm 

long, subglabrous to sparsely ciliate. Tepals unequal (outer tepals shorter by c. 1.5 mm), 

glabrous, lanceolate, 4.2–5.2 (–5.5) mm long, snow-white, becoming slightly pale strawwhite, 

or rarely suffused purplish. Stamens 6, longer than outer tepals; anthers 1.8–2.2 mm 

long, turning brownish after dehiscence; filaments 1.5–1.9 mm long; style 2.2–2.6 mm long; 

stigmas 1.0–1.3 mm long. Capsule brown, shining, acuminate and subtrigonous above, 

mucronate; capsule segments usually 2.0–2.4 mm long including a mucro c. 0.5 mm long and 

c. 1.5 mm wide. Seeds ellipsoidal, dark castaneous-brown, 1.2–1.4 mm long including a 

distinct apical protuberance 0.2–0.3 mm long and c. 0.7 mm wide; appendage yellowish, 

c. 0.1 mm long, fibrillate. 2n=12, fide H.Nordenskiöld, Hereditas 37: 328–329 (1951); 

Druskovic, Int. Organ. Pl. Biosyst. Newslett. 24: 14 (1995). 

Found in the Alps and Pyrenees, from Austria, Slovenia and Germany to NE Spain and S and 

C France. It is commonly cultivated and may escape from gardens. 11: AUT, GER, SWI. 

12: FRA, SPA. 13: ITA, YUC-SL. Open patches in mountain woodlands and near the 

timberline, usually on calcareous soils. Map 22. 

11. AUSTRIA: Tirol, Gschnitz, Trins, 1300 m, F.Kerner [Fl. Exs. Austro-Hung.] 1870 (PR, WU). 

GERMANY: Oberbayern, Loisach-Auen, Scharnitz, D.Podlech 34413 (LD, M); Oberbayern, Mittenwald, 

930 m, F.Vollmann [Fl. Exs. Bavar.] 1363 (M, PR). SWITZERLAND: Valais, Orcières, 1480 m, J.Miège et 

al. [Exs. Genav.] 196 (W). 12. FRANCE: Puy-de-Dôme, Mont-Dore, Capucin, 1150 m, 4 Aug 1883, 

P.Billiet [C.Magnier, Fl. Select. Exs.] 987 (W, LD). SPAIN: Huesca, Torla, Turieto Bajo, 1200 m, 

P.Montserrat 4745/70 (JACA, LD); Aragón, Puerto de Búcaro, Bordère (BC, PR). 13. ITALY: Valle d' 

Aosta, Grand St. Bernard, 2000–2500 m, R.K.Brummitt 5447 (K); Lombardia, Corno, Mt Grigna, 1800 m, 

1895, J.M.Norman (S). SLOVENIA: Julijske Alpe, Bohinjsko jezero, 800 m, F.Dolšak (PR); Kranjska 

Gora, Veršec, 18 Jul 1909, J.Paul 119 (PR). 

25

26 


7. Luzula pedemontana Boiss. & Reut., Pugill. Pl. Afr. Bot. Hispan. 115 (1852) 

Juncoides pedemontana (Boiss. & Reut.) Kuntze, Revis. Gen. Pl. 2: 725 (1891); Luzula nivea subsp. 

pedemontana (Boiss. & Reut.) Bonnier & Layens, Tabl. Syn. Pl. Vasc. France 322 (1894). T: Italy, 

Piemonte, Col de Tende, P.E.Boissier & G.F.Reuter; syn: G, n.v.; Col de Tende, H.G.Reichenbach [Fl. Germ. 

Exs.] 2315; syn: K. 

Luzula nemorosa var. gracilis E.Mey., Linnaea 22: 395 (1849). T: Italy, Piemonte, Col de Tende, 

H.G.Reichenbach [Fl. Germ. Exs.] 2315; syn: K; O.W.Sonder; syn: n.v. 

Illustrations: F.G.P.Buchenau, in H.G.A.Engler, Pflanzenr. (iv.36) 25: 52, fig. 41 (1906); H.E.Hess, E.Landolt 

& R.Hirzel, Fl. Schweiz 1: 517 (1967). 


Perennials 25–50 cm tall, loosely caespitose, ciliate; rhizome thin, creeping to ascending; 

stolons ascending, with base covered with brown cataphylls. Basal leaves usually 8–10 (–12) cm 

long, flat to involute, up to 2.5 (–3.5) mm wide; leaves of young shoots very narrow, 

involute; cauline leaves 3–5, of variable length, up to 3.5–4.0 mm wide but often very 

narrow, c. 1 mm wide. All leaves distinctly acuminate; margins papillose-serrate. Lower 

bract subherbaceous, up to 4–5 cm long, shorter than inflorescence. Inflorescence 

decompound, lax, erect to ±nodding, corymbiform; main branches often arcuate, each with 

loose anthela of 2–7-flowered clusters. Bracteoles whitish-membranous, ovate-lanceolate, 

c. 2.0–2.5 mm long, sparsely ciliate. Tepals ±equal, oblong-lanceolate, glabrous, 3.1–3.6 mm 

long, white or dirty white to whitish-membranous. Stamens 6; anthers 1.3–1.6 mm long, 

upper part (up to one third) bifurcate; filaments 0.4–0.6 mm long; style 1.4–1.9 mm long; 

stigmas 0.6–1.0 mm long. Capsule oblong-ovoid, long acuminate, mucronate, castaneousbrown, 

shining; capsule segments (2.3–) 2.4–2.7 (–2.9) mm long, including a mucro 

c. 0.2–0.4 mm long and 1.5–1.6 mm wide. Seeds ellipsoidal, dark brown, 1.3–1.4 mm long 

including a pale apex 0.1–0.2 mm long and c. 0.8 mm wide; basal appendage c. 0.1 mm long, 

fibrillate. 

Restricted to SE France, N Italy and Corse. 12: COR, FRA. 13: ITA. Open woodlands and 

scrub in foothills and mountains. Map 23. 

12. CORSE: Foret d'Aitone, E.Reverchon [Pl. Corse] 443 (PR, UPS, W); Vizzavona, 24 Jun 1914, K.Ronniger 

(W); Restonica, 1350 m, H.Malicky 9585 (W). FRANCE: Alpes Maritimes, Muntis sur Fontan, E.Reverchon 

[Pl. Franc.] 128 (K, LD, UPS); Alpes Maritimes, Col di Tenda, 31 Jul 1879, J.Vetter (K). 13. ITALY: 

Liguria, San Remo, Bajardo, C.Bicknell [I.Dörfler, Herb. Norm.] 3268 (PR, W); Toscana, Alpi Apuanae, Mt 

Procinto, 600 m, S.Snogerup 1603 (LD); Piemonte, Piossano, Prese di Gangano, 27 Jun 1926, P.Fontana & 

F.Santi (UPS); Piemonte, Cuneo, Monticello d'Alba, 300 m, T.Ferraris & G.Ferro [Fl. Ital. Exs.] 513 (K); 

Piemonte, Val Pesio, San Bartolomeo, 750 m, C.Pollini [A.Kneucker, Cyper. Junc. Exs.] 22 (PR). 

8. Luzula lutea (All.) DC., in J.B.A.P. de M. de Lamarck & A.P. de Candolle, Fl. 

Franç., 3rd edn, 3: 159 (1805) 

Juncus luteus All., Fl. Pedem. 2: 216 (1785); Juncoides lutea (All.) Kuntze, Revis. Gen. Pl. 2: 724 (1891); 

Juncus multiflorus Retz., Fl. Scand. Prodr., 2nd edn, 82 (1795). T: Haller 1329 [V.A. von Haller, Hist. 

Stirp. Helv. 2: no. 1329 (1768)]; syn: P-Haller; A. von Haller, Agrostographia, no. 314 (1775) [Switzerland] 

in montibus Rhaeticis, speciatim in Aviculae montis summis; syn: n.v.; [authentic material: K]. 

Juncus spadiceus Vill., Prosp. Hist. Pl. Dauphiné 18 (1779). T: ‘Hall. Enum. 257. n. 18’; syn: P-Haller, 

n.v. [Luzula spadicea [All.] DC. is not based on the same type.] 

Juncus aureus Pourr., Hist. & Mém. Acad. Roy. Sci. Toulouse 1: 321 (1786), n.v.; Luzula aurea (Pourr.) 

Timb.-Lagr., Bull. Soc. Sci. Phys. Nat. Toulouse 2: 132 (1875), n.v. T: ‘A. Llaurenti’, Pourret; syn: UPS- 

Thunberg; Haller 1329 [Haller, Hist. Stirp. Helv. 2: no. 1329 (1768)]; syn: P-Haller. 

Luzula lutea (All.) DC. subsp. pyrenaica P.Monts., Anales Inst. Bot. Cavanilles 21(2): 438 (1964). 

T: Spain, Pyrenees, Estanys d'En Gaït, Circ d'En Gaït, 2500 m, Aug 1949, P.Montserrat; holo: BCF. 

Illustration: H.E.Hess, E.Landolt & R.Hirzel, Fl. Schweiz 1: 516 (1967). 


Perennials (10–) 12–15 (–25) cm tall, loosely caespitose, ±glaucous-green, subglabrous; 

rhizome creeping to ascending; stolons ascending, covered with purplish brown cataphylls or


with sparse fibrous cover. Basal leaves usually (3–) 4–7 (–9) cm long, (3–) 4–5 (–6) mm 

wide; cauline leaves usually 3, usually 2–4 (–6) cm long. All leaves subacute to subobtuse; 

margins distinctly papillose-serrulate. Lower bract ±brownish-membranous, 1.0–1.5 (–2.0) cm 

long, shorter than inflorescence. Inflorescence decompound, corymbiform; main branches 

thin, straight to arcuate; lower branches often patent, sometimes erect, each with dense 

glomeruliform (5) 10–15 (25)–flowered cluster, with one cluster often larger, subsessile. 

Bracteoles ovate, c. 1.5 mm long, transparent-scarious, ±ciliate. Tepals unequal, ±oblonglanceolate, 

glabrous, straw-yellow, becoming ±dirty yellow, with broad membranous 

margins; inner tepals 2.8–3.2 mm long; outer tepals shorter, c. 2.5–2.7 mm long. Stamens 6; 

anthers 1.1–1.8 mm long; filaments 0.3–1.1 mm long, distinctly elongating during flowering 

(anther/filament ratio thus changing); style (0.9–) 1.0–1.3 mm long; stigmas 0.5–0.8 mm 

long. Capsule oblong-ovoid, acuminate, mucronate, deep castaneous-brown, shining; capsule 

segments 2.4–2.8 × 1.4–1.5 mm long including a mucro 0.2–0.3 mm long. Seeds oblongovoid, 

paler brown, 1.0–1.3 mm long including a paler apex c. 0.2 mm long and 0.7–0.8 mm 

wide; appendage c. 0.1 mm, fibrillate. 2n=12, fide H.Nordenskiöld, Hereditas 37: 328–329 

(1951). 

Southwestern part of the Alps, E Pyrenees and N Italian mountains. 11: AUT, SWI. 12: FRA, 

SPA. 13: ITA. Growing in alpine grasslands and screes, usually on siliceous substrates, from 

c. 1700 m. Map 24. 

11. AUSTRIA: Nordtirol, Gurgler Tal, 1900–2200 m, 13 Sep 1989, A.Polatschek 17612 (W); Oetztal, 

Klebealpe, 2000 m, 11 Jun 1937, Fischer (K). SWITZERLAND: Valais, Saas–Fée, 2100 m, L.Marret [Exs. 

Fl. Valais] 296 (S); Valais, Alpe de la Barma, 2200 m, F.O.Wolf [A.Kneucker, Cyper. Junc. Exs.] 21 (PR); 

Graubünden, Ober Engadin, Bernina Pass, 2330 m, O.Hedberg 4022 (UPS). 12. FRANCE: Haute-Savoie, 

Massif du Mont Buet, Val de Bérano, 2000 m, B. de Retz 40427 (W); Hautes-Alpes, Goudran, Montgenèvre, 

2000–2200 m, C.Magnier [Fl. Select. Exs.] 1823 (LD). ANDORRA: Tristaina Lakes, 2800 m, Lewis 485 (K). 

SPAIN: Setcasas, Mt Morens, E.Bourgeau [Pyren. Esp.] 277 (K). 13. ITALY: Levico, Pancrotto, Jul 1926, 

Meebold (K). 

A species with limited variation. The westernmost populations were recognized as subsp. 

pyrenaica on the basis of the inflorescence habit, broader leaves, longer tepals and smaller 

seeds. However, despite the fact that eastern populations show a lower frequency of broadleaved 

individuals, the other characters do not exhibit any geographical tendency, and the 

extremes are often found within populations throughout the geographical range of the species. 

9. Luzula luzuloides (Lam.) Dandy & E.Willm., J. Bot. 76: 352 (1938) 

Juncus luzuloides Lam., Encycl. 3: 272 (1789). T: France: ‘Lorraine Allemande, près des rives de la Sarre’, 

de l’Aumont; lecto: P-LAM, fide S.Snogerup, in P.H.Davis, Fl. Turkey 9: 28 (1986). 

Luzula albida var. macrantha Beck, Fl. Nieder-Österreich 1: 159 (1890); Luzula nemorosa A.I. [unranked] 

macrantha (Beck) Asch. & Graebn., Syn. Mitteleur. Fl. 2(2): 502 (1904); Luzula albida subvar. macrantha 

(Beck) Rouy, Fl. France 13: 261 (1912). T: Austria, Mt Schneeberg, G.Beck; holo: n.v. 

Luzula albida var. parviflora Döll, Fl. Baden 1: 323 (1857); Luzula nemorosa var. parviflora (Döll) Asch. & 

Graebn., Syn. Mitteleur. Fl. 2(2): 504 (1904); Luzula luzuloides var. parviflora (Döll) I.Grint., Fl. Rep. Soc. 

Roman. 11: 591 (1966). T: [Germany] Baden, Untergrombach, Döll; syn: n.v. 

Luzula nemorosa B.II. [unranked] hylogiton Asch. & Graebn., Syn. Mitteleur. Fl. 2(2): 504 (1904). 

T: Germany, Harz; n.v. 

Maps: H.Meusel, E.Jäger & E.Weinert, Vergl. Chorol. Zentraleur. Fl. 1: 87 (1965); E.Hultén & M.Fries, Atlas 

N. Eur. Vasc. Pl. 1: map 165 (1986). 

Perennials usually 35–75 cm tall, loosely caespitose to caespitose, densely ciliate; rhizome 

horizontal to ascending, branched; stolons usually present, ascending. Basal leaves 

numerous, usually 12–18 (–30) cm long, width variable but ranging from (3–) 4–5 (–6) mm; 

cauline leaves usually 4–5, to c. 25 cm long, with the upper ones often overtopping 

inflorescence. All leaves with acute tip; margins papillose-serrulate. Lower bracts herbaceous, 

of variable length, usually overtopping inflorescence, less often shorter. Inflorescence rich, 

decompound, ±anthelate; main branches usually numerous, with secondary branches each 

bearing ±dense or lax anthela of (2–) 3–6 (–8)-flowered clusters. Bracteoles ±ovate, 

subglabrous to ciliate, usually 1.0–1.5 mm long. Tepals glabrous, subequal with inner longer 

by c. 0.5 mm, ±lanceolate, (2.0–) 2.3–3.8 (–4.2) mm long. Stamens 6; anthers 1.0–1.8 mm 

27

28 


long; filaments 0.3–0.6 mm long; style 0.8–1.2 mm long; stigmas 0.6–c. 1.1 mm long. 

Capsule oblong-ovoid, gradually acuminate, mucronate, ±castaneous-brown, ±shining; 

capsule segments 2.2–2.6 mm long including mucro 0.2–0.5 mm long and 1.1–1.3 mm wide. 

Seeds ellipsoidal, ±dark brown, usually 1.2–1.4 mm long including apical protuberance 

c. 0.2 mm long and 0.7–0.8 mm wide; appendage indistinct, c. 0.1 mm, fibrillate. 

Europe, from the Netherlands, France and N Spain throughout C Europe to Greece, Turkeyin-Europe 

and W Russia. Introduced to N and E Europe and E North America, often 

naturalized. There are two variants, from lowland and mountain areas; the two extremes are 

locally linked by numerous intermediates. 

Tepals ±white, dirty white or ±scarious, usually (2.0–) 2.3–3.2 mm long 

Tepals usually coppery-coloured, brownish-purplish or pale castaneous with 

9a. subsp. luzuloides 

reddish tinge, usually 3.0–3.5 (–4.2) mm long 9b. subsp. rubella 

9a. Luzula luzuloides (Lam.) Dandy & E.Willm. subsp. luzuloides 

Juncus nemorosus Pollich, Hist. Pl. Palat. 1: 352 (1776); Juncus luzula Krock., Fl. Siles. 5(1): 569 (1787), 

nom. illeg.; Juncus maximus var. angustifolius Roth, Tent. Fl. Germ. 1: 157 (1788); Juncus angustifolius 

Wulfen, in N.J. von Jacquin, Collectanea 3: 56 (1789), nom. illeg.; Juncus leucophobus Ehrh., Beitr. Naturk. 

6: 141 (1791), nom. illeg.; Luzula angustifolia [Wulfen] Wender., Beitr. Fl. Hess. 36 (1823), nom. illeg., non 

Poir.; Luzula nemorosa (Pollich) E.Mey., Linnaea 22: 394 (1849), nom. illeg., non Hornem. (1815), nec 

Baumg. (1816); Juncoides nemorosa (Pollich) Kuntze, Revis. Gen. Pl. 2: 724 (1891). T: Germany, 'circa 

Lauteren', Pollich. 

Juncus albidus Hoffm., Deutschl. Fl. 126 (1791); Luzula albida (Hoffm.) DC., in J.B.A.P. de M. de Lamarck 

& A.P. de Candolle, Fl. Franç., 3rd edn, 3: 159 (1805); Luzula nivea subsp. albida (Hoffm.) Bonnier & 

Layens, Tabl. Syn. Pl. Vasc. France 322 (1894). T: icon, in G.F.Hoffmann, Deutschl. Fl., fig. 4 (1791); syn; 

icon, in J.D.Leers, Fl. Herborn., Pl. 13, fig. 9 (1775) [ut J. niveus]; syn. 

Luzula albida var. leucanthema Wallr., Sched. Crit. 1: 150 (1822); Luzula nemorosa A.II.a. [unranked] 

leucanthema (Wallr.) Asch. & Graebn., Syn. Mitteleur. Fl. 2(2): 502 (1904). T: [Germany, Halle], 

C.F.W.Wallroth; syn: PR; C.Schkuhr, Bot. Handb. 1: plate 98 (1791); syn. 

Luzula nemorosa A.II.a.2. [unranked] laxa Asch. & Graebn., Syn. Mitteleur. Fl. 2(2): 503 (1904). T: not 

designated. 

Illustrations: T.Szynal & J.Mądalski, Atlas Fl. Polsk. 2(2): 152a (1931); H.E.Hess, E.Landolt & R.Hirzel, 

Fl. Schweiz 1: 517 (1967). 

Plants usually very loosely caespitose; stoloniferous. Bracteoles membranous-scarious. 

Tepals ±white, dirty white or ±scarious, usually (2.0–) 2.3–3.2 mm long. 2n=12, fide 

H.Nordenskiöld, Hereditas 37: 328–329 (1951). Fig. 6. 

Most common in C Europe; otherwise throughout the range of the species. Introduced to and 

often naturalized in N and E Europe and E North America. 10: DEN, fin, grb, nor, SWE. 

11: AUT, BGM, CZE, GER, HUN, NET, POL, SWI. 12: FRA, SPA. 13: ALB, BUL, GRC, 

ITA, ROM, TUE, YUC. 14: BLR, BLT, ruc, run, ruw, UKR. 72: nsc, ont. 74: min, wis. 75: 

cnt, mai, mas, nwy, pen. Found in broad-leaved woodlands, clearings, less often meadows, 

from lowlands to foothills. Map. 25 [only European introductions shown, not distinguished 

from the native occurrence]. 

10. FINLAND: Turku, Ruissalo, 13 Jul 1923, V.Pesola (PR 76409); Turku, Ruissalo, S. Valle [Pl. Finl. Exs.] 

1862 (H, K, WU). 11. AUSTRIA: Karawanken, Freiberg, 28 Jul 1970, Pull (W); N Tirol, between Kössen 

and Klobenstein, 590 m, 16 Jun 1972, A.Polatschek (W); Wienerwald, Königstätten, Gölles 266 (W). 

GERMANY: Sachsen, Pappritz, 23 May 1904, C.Marquardt (PR); Baden, Karlsruhe, Durlacher Wald, 117 m, 

A.Kneucker [Cyper. Junc. Exs.] 23 (PR, W). 12. FRANCE: Ardennes, Bois de Chagny, Mézières, E.Callay 

[Puel & Maille, Fl. Région. Fr.] 79 (W); Vosges, between Igney and Nomexy, Gérard [C.Magnier, Fl. Select. 

Exs.] 2304bis (LD). 13. ALBANIA: Skutari, Mt Zukali, A.Baldacci [It. Alban, 5] 47 (WU). ITALY: 

Padova, Teolo, Mt Pendice, 200 m, A.Fiori & A.Béguinot [Fl. Ital. Exs.] 2015 (K, WU). 14. RUSSIA 

NORTHWEST: Vyborg, Monrepos, K.Nilsson 30/38 (H). RUSSIA NORTH: Sortavala, 24 Jun 1921, 

K.H.Hällström (H).


Figure 6. Luzula luzuloides subsp. luzuloides. A, habit; B, terminal part of inflorescence; 

C, flower; D, capsule; E, F, tepals; G, open capsule; H, seed; I, stem T.S.; J, leaf T.S. 

(A–C, E-G, I, J, Stelzer s.n., 1883, KRA; D, H, J.M ądalski s.n., 13 Aug 1928, KRA). Scale 

bars: A = 5 cm; B = 2 mm; C–H = 2 mm; I, H = 1 mm. Drawn by T.Szynal. Reproduced 

with permission, from T.Szynal & J.Mądalski, Atlas Fl. Pol. 2/2: 152a (1931). 

29

30 


9b. Luzula luzuloides subsp. rubella (Hoppe ex Mert. & W.D.J.Koch) Holub, Folia 

Geobot. Phytotax. 18: 205 (1983) 

Luzula albida var. versicolor Bluff & Fingerh., Comp. Fl. German. 1: 443 (1825); Luzula albida var. rubella 

Hoppe ex Mert. & W.D.J.Koch, in Deutschl. Fl.(Röhling) 2: 599 (1826); Luzula nemorosa var. rubella (Hoppe 

ex Mert. & W.D.J.Koch) Garcke, Fl. N. Mitt.-Deutschland, 2nd edn, 340 (1851); Luzula angustifolia var. 

rubella (Hoppe ex Mert. & W.D.J.Koch) Garcke, Fl. N. Mitt.-Deutschland, 3rd edn, 348 (1853); Luzula 

nemorosa f. rubella (Hoppe ex Mert. & W.D.J.Koch) Racib., Consp. Juncac. Polon. 25 [p. separ.] (1888); 

Luzula angustifolia subsp. rubella (Hoppe ex Mert. & W.D.J.Koch) K.Richt., Pl. Eur. 1: 184 (1890); Luzula 

albida subvar. rubella (Hoppe ex Mert. & W.D.J.Koch) Rouy, Fl. France 13: 261 (1912). T: Austria, 

Heiligenblut, D.H.Hoppe [Herb. Viv. Pl. Gram.] 68; syn: LI, BRNM etc. (further authentic material: K, LD, 

PR). 

Luzula albida var. cuprina Rochel, Pl. Banat. Rar. 3: 27 (1828), nom. inval.; Luzula nemorosa A.II.b. 

[unranked] cuprina [Rochel ex] Asch. & Graebn., Syn. Mitteleur. Fl. 2(2): 503 (1904); Luzula luzuloides 

subsp. cuprina [Rochel ex] Chrtek & Křísa, Preslia 46: 212 (1974). T: Slovakia, 'ad pedes Kralowahora', 

A.Rochel; authentic material: W, PR [as Juncus albidus var. cuprinus]. 

Luzula nemorosa var. colorata E.Mey., Linnaea 22: 395 (1849). T: in Alpibus. D.H.Hoppe; syn: LI; in 

Carpathis, Baumgarten; syn: n.v. 

Luzula alpigena Schur, Enum. Pl. Transsilv. 681 (1866). T: n.v. [numerous syntypes from Romanian South 

Carpathians, most probably in LW]. 

Luzula albida var. erythranthema Wallr., Sched. Crit. 1: 150. (1822); Luzula nemorosa var. erytranthema 

(Wallr.) Hayek, Fl. Stir. Exs., no. 409 (c. 1930) [printed exsiccatum label]; Luzula luzuloides var. 

erythranthema (Wallr.) I.Grint., Fl. Rep. Soc. Roman. 11: 592 (1966). T: Austria, Heiligenblut, D.H.Hoppe; 

syn: PR [specimen annotated by C.F.W.Wallroth]; Heiligenblut, D.H.Hoppe [Herb. Viv. Pl. Gram.] 68; syn: 

LI, BRNM. 

Luzula fuscata Schur, Verh. Mitth. Siebenbürg. Vereins Naturwiss. Hermannstadt 10: 200 (1859). 

T: [Romania], 'Am Büdöskegel', F.Schur; holo, n.v. [doubtfully valid]. 

Luzula nemorosa [unranked] fuliginosa Asch., Fl. Brandenburg 1: 740 (1864); Luzula albida var. fuliginosa 

(Asch.) Beck, Fl. Nieder-Österreich 1: 159 (1890). T: [Czech Republic, the Krkonoše, Labský vodopád], 

am Elbfall des Riesengebirges, Jul 1862, C.Lucas; holo: n.v. 

Luzula nemorosa f. rosea Kožuharov, Izv. Bot. Inst. (Sofia) 11: 127 (1963), nom. inval. 

Luzula nemorosa f. grandiflora Kožuharov, Izv. Bot. Inst. (Sofia) 11: 127 (1963), nom. inval., & in 

D.Jordanov, Fl. N. R. B'lgar. 2: 401 (1964), nom. inval. T: Bulgaria, Mt Balkan centralis [collector not 

given]; holo: n.v. 

Luzula nemorosa f. aggregata Kožuharov, Izv. Bot. Inst. (Sofia) 11: 127 (1963). T: Bulgaria, Vitosha, 

A.Yavashev; holo: SOM. 

Luzula luzuloides subsp. tenacissima Vivant, Bull. Soc. Bot. France 126: 259 (1979). T: France, 

W Pyrenees, Haute-Soule, Pic Lackhoura, 1700–1800 m, 5 Sep 1973, J.Vivant s.n.; holo: P; iso: JACA. 

Illustration: T.Szynal & J.Mądalski, Atlas Fl. Polsk. 2(2): 152 (1931). 

Plants loosely to ±densely caespitose; stolons short, ascending. Bracteoles usually pale 

castaneous or coppery-coloured. Tepals usually coppery-coloured, brownish-purplish or pale 

castaneous with reddish tinge, usually 3.0–3.5 (–4.2) mm. 2n=12, fide C.Garcia-Herran, in 

litt. (material from Polish Tatra Mtns and Bieszczady, Poland, vouchers in NEU). Fig. 7. 

Mountains of C and S Europe, the Pyrenees, introduced to Great Britain. 10: grb. 11: AUT, 

CZE, GER, POL, SWI. 12: FRA, SPA. 13: ALB, BUL, GRC, ITA, ROM, YUC. 14: UKR. 

Mountain to subalpine tall-grass meadows, open mountain woodlands; in mountain regions 

where it is common, it is also found in valley situations. Map 26. 

11. AUSTRIA: Oetztaler Alpen, Landeck, Venetberg, 2300 m, Aug 1968, A.Polatschek (W); Steiermark, 

Pretulalpe, Fischbacher Alm, 16 Jul 1911, K.Ronniger (W). CZECH REPUBLIC: Hrubý Jeseník, Petrovy 

kameny, 27 Jul 1889, V.Spitzner (PR 49587). POLAND: Babia Góra, 1480 m, Walas [Pl. Polon. Exs.] 81 (K). 

13. BULGARIA: Vitoša, J.Velenovský (PR 49584). CRNA GORA: Sekirica Planina, Jul 1903, J.Rohlena (PR). 

ITALY: South Tyrol, Dolomiti, Villnoss, Zamseralpe, 1912, K.Ronniger (W). F.Y.R. MACEDONIA: Skopje, 

Šar Planina, Ljuboten, 12 Jul 1959, J.D.A.Stainton 7988 (K). 

Variable in leaf width, size of inflorescence and perianth colour. Intermediates between 

subspecies are not rare, e.g., 11. POLAND: Nowy Targ, Pieniny, Stolarzówka, 620–640 m, 

H.Piękoś [Fl. Polon. Exs.] 389 (KRA, KRAM, W). Maybe better treated as a variety.


Figure 7. Luzula luzuloides subsp. rubella. A, habit; B, terminal part of inflorescence; 

C, H, flowers; D, capsule; E, seed; F, G, tepals; I, stem T.S.; J, leaf T.S. 

(A–C, F–I, A.Wołoszczak, Aug 1887, KRA; D, E, J.M ądalski s.n., 10 Aug 1927, KRA). 

Scale bars: A = 5 cm; B = 5 mm; C–H = 2 mm; I, J = 1 mm. Drawn by T.Szynal. 

Reproduced with permission, from T.Szynal & J.Mądalski, Atlas Fl. Pol. 2/2: 152 (1931). 

31

32 


10. Luzula sylvatica (Huds.) Gaudin, Agrost. Helv. 2: 240 (1811) 

Juncus sylvaticus Huds., Fl. Angl. 32 (1762); Juncoides sylvatica (Huds.) Druce, Fl. Oxfordshire 434 (1927). 

T: England, Hamstead, Hudson; syn: not extant; Juncus foliis planis latissimis...Hall. helv. 257 [A. von Haller, 

Enum. Meth. Stirp. Helv. Ind. 1: 257 (1742)]; [Switzerland; citation from Haller (1742): in sylvis opacis: v.g. 

montis Falconarii, circa Tigurum Scheuchzer]; syn: P-Haller. 

Luzula haussknechtiana Freyn & Sint., Bull. Herb. Boissier, ser. 1, 4: 200, 1896. T: Turkey, Darsos Dagh, 

P.E.E.Sintenis 5775; syn: BM, BRNM, PR, WU. 

Luzula sylvatica subsp. croatica Beyer, Verh. Bot. Vereins Prov. Brandenburg 41: xix (1900); Luzula 

sylvatica var. croatica (Beyer) Buchenau, in H.G.A.Engler, Pflanzenr. (iv.36) 25: 55 (1906). T: Croatia, 

‘Klek bei Ogulin’, Musulinski Potok, R.Beyer; syn: n.v.; Croatia, 'Turcovic Sello', R.Beyer; syn: n.v. 

Luzula sylvatica var. dertosensis P. Monts., Anales Inst. Bot. Cavanilles 21(2) [1963]: 441 (1964). 

T: Spain, Ports de Tortosa, Font dels Bassis, 750 m, 25 May 1917, P.Font Quer; holo: BC; iso: BC. 

Luzula sylvatica A.I.a.b.[unranked] pulchra Asch. & Graebn., Syn. Mitteleur. Fl. 2(2): 499 (1904). T: SW Alps 

[otherwise not indicated]. 

Luzula sylvatica f. altissima Kožuharov, Izv. Bot. Inst. (Sofia) 11: 126 (1963), nom. inval. 


N. Eur. Vasc. Pl. 1: map 164 (1986). 

Perennials, usually up to 100 cm tall, caespitose to loosely caespitose; rhizome stout, long, 

branched, horizontal to ascending; stolons ±absent. Basal leaves up to 40 cm long, usually 

4–12 mm wide; cauline leaves 3–5 (–7); leaf tip ±acute; margins ±smooth. Lower bract 

(1.2–) 2.0–3.0 (–5.0) cm long, shorter than inflorescence. Inflorescence decompound, usually 

lax, up to 20 cm long; secondary anthelas with 5–45 flowers; terminal clusters (1–) 2–3 (–5)flowered. 

Bracteoles whitish-membranous, ovate, up to c. 2 mm long, sparsely ciliate. Tepals 

subequal; outer tepals usually 2–4 (–5) mm long, oblong-lanceolate, pale castaneous-brown 

to deep brown; margins usually hyaline. Stamens 6; anthers 1.0–3.7 mm long; filaments 

usually 0.3–0.7 mm long; style 1.6–2.2 mm long; stigmas 2.2–2.8 mm long. Capsule 

castaneous-brown to dark brown, ±oblong-ovoid, ±gradually narrowed, usually 2.0–4.0 mm 

long including mucro 0.2–0.8 mm long. Seeds brown to dark brown, oblong-ovoid, 1.2–1.9 mm 

long, usually 0.7–1.2 mm wide; appendage 0.1–0.2 mm, fibrillate. 

Found in Europe, except for most of Scandinavia and Russia; outside Europe only marginally 

in the W Caucasus, Transcaucasia and N Turkey. A very variable species, particularly in the 

southern part of its range. Several subspecies have been described; some other aberrant 

populations remain to be studied, all in the S European peninsulas. Four subspecies are 

recognized here. In addition, some (not all) plants and populations from the Balkan peninsula, 

most typically from Greece, differ from the typical L. sylvatica in a number of characters, 

e.g., in having basal leaves (6–) 8–10 (–11) mm wide, inflorescence lax, few-flowered, 

secondary anthelas usually with 8–14 (–16) flowers, terminal clusters (1–) 2–4 (–5)flowered, 

tepals up to 4 (–5) mm long, capsule 3.1–3.9 mm long and seeds up to 1.2 mm 

wide (representative material at LD). They may represent another intraspecific taxon of 

L. sylvatica; many intermediates between these aberrants and more common forms of subsp. 

sylvatica are known from other parts of the Balkans and from Turkey. The type subspecies is 

often cultivated as an ornamental plant. 

1 Tepals usually up to 2.7 mm long; primary branches of inflorescence 

usually elongated; secondary anthelas usually ±contracted, rarely ±lax 10b. subsp. henriquesii 

1: Tepals usually more than 2.7 mm long; primary branches of inflorescence 

usually not distinctly elongated, ±evenly branched to form ±lax 

secondary anthelas 

2 Basal leaves with margins and base very densely long ciliate; tepals 

longer than 4 mm; secondary anthelas usually with up to 15 flowers; 

anthers (1.7–) 2.2–2.5 (–3.0) mm long; leaf tip elongated, acuminate; 

seeds 1.1 mm wide or wider 10c. subsp. sicula 

2: Basal leaves with margins ciliate but not conspicuously densely or long; 

tepals up to 3.7 mm long; anthers up to 1.9 mm long; leaf tip acute to 

subobtuse-mucronate; seeds up to 0.8 mm wide


3 Plants usually 50–100 cm tall; basal leaves (7–) 9–12 mm wide; 

capsule usually equalling perianth 10a. subsp. sylvatica 

3: Plants usually up to 55 (–70) cm; basal leaves usually 5–8 mm wide; 

capsule equalling or shorter than perianth 10d. subsp. sieberi 

10a. Luzula sylvatica (Huds.) Gaudin subsp. sylvatica 

Juncus maximus Reichard, Fl. Moeno-Francof. 182 (1772); Juncus pilosus subsp. maximus (Reichard) Ehrh., 

Hannover. Mag. 1782(23): 363 (1782); Luzula maxima (Reichard) DC., in J.B.A.P. de M. de Lamarck & 

A.P. de Candolle, Fl. Franç., 3rd edn, 3: 161 (1805). T: Haller 1324 [A. von Haller, Hist. Stirp. Helv. 

2: no. 1324 (1768)]; syn: P-Haller; 'ad rivulum an der Reichenbach'; syn: n.v.; 'bey den drey Born nicht weit 

von Falckenstein'; syn: n.v. 

Juncus latifolius Wulfen, in N.J. von Jacquin, Collectanea 3: 59 (1789); Luzula maxima var. latifolia 

(Wulfen) Čelak., Prodr. Fl. Böhmen 1: 84 (1867), nom. inval.; Luzula sylvatica var. latifolia (Wulfen) 

Buchenau, in H.G.A.Engler, Pflanzenr. (iv.36) 25: 55 (1906). T: ‘in superioribus umbrosis sylvis alpium 

Lithopolitanarum’, F.X.Wulfen; syn: probably destroyed; Haller 1324 [A. von Haller, Hist. Stirp. Helv. 2: 

no. 1324 (1768)]; syn: P-Haller. 

Juncus montanus Lam., Encycl. 3: 273 (1789). T: France, Mont d'Or, J.B.A.P.Lamarck; syn: P-LAM; Haller 

1324 [A. von Haller, Hist. Stirp. Helv. 2: no. 1324 (1768)]; syn: P-Haller. 

Juncus pilosus L. A. [unranked] fascicularis Schrank, Baier. Fl. 1: 621 (1789). T: [Germany] Bayern 

[otherwise not indicated]; Haller 1324 [A. von Haller, Hist. Stirp. Helv. 2: no. 1324 (1768)]; syn: P-Haller. 

Luzula sylvatica var. gracilis Rostr. ex Beeby, Scott. Nat. 1887: 29 (1887), n.v. [Also in Buchenau, Bot. Jahrb. 

Syst. 12: 92. 1890]; Luzula maxima var. gracilis (Rostr. ex Beeby) Husn., Joncées 22 (1908). T: Faeroes, on 

the top of Hestoe, E.Rostrup; syn: C, n.v.. 

Luzula sylvatica A.I.a.2. [unranked] pallida Asch. & Graebn., Syn. Mitteleur. Fl. 2(2): 499 (1904). T: [not 

indicated]. 

Illustrations: T.Szynal & J.Mądalski, Atlas Fl. Polsk. 2(2): 153 (1931); H.E.Hess, E.Landolt & R.Hirzel, Fl. 

Schweiz 1: 519 (1967). 

Plants usually 45–100 cm tall. Basal leaves usually (7–) 9–12 (–18) mm wide, gradually to 

abruptly narrowed into acute tip; margins ciliate. Inflorescence usually lax; secondary 

anthelas usually with 15–35 flowers; terminal clusters (1–) 2–3 (–4)-flowered. Outer tepals 

usually (2.1–) 2.7–3.0 (–3.7) mm long. Anthers usually 1.0–1.9 mm long; filaments 

(0.2–) 0.3–0.5 (–0.7) mm long; style 1.6–2.0 mm long; stigmas 2.5–2.8 mm long. Capsule 

usually ±equalling perianth, oblong-ovoid, (1.8–) 2.5–2.8 (–3.4) mm long, slightly narrowing 

to a mucro usually 0.2–0.5 mm long. Seeds (1.0–) 1.3–1.5 (–1.8) mm long, up to 0.8 mm 

wide; appendage c. 0.1 mm long. 2n=12, fide C.Garcia-Herran (see list of specimens); 

H.Nordenskiöld, Hereditas 37: 328–329 (1951). Fig. 8. 

West and C Europe, from Lofoten, Faeroes, SW Norway, Great Britain and most of S Europe 

to the Carpathians, N Turkey and the Caucasus. 10: DEN, FOR, GRB, IRE, NOR, swe. 

11: AUT, BGM, CZE, GER, NET, POL, SWI. 12: COR, FRA. 13: ALB, BUL, GRC, ITA, 

ROM, YUC. 14: RUS?, UKR. 33: NCS, TCS. 34: TUR. Woodlands from lowlands to the 

mountains, or damp grasslands, open moorland and along streams. Map 27. 

10. DENMARK: Vejle, Munkebjerg, K.Larsen & P.Pedersen [Fl. Jutl. Exs.] 4 (AAU, PR). GREAT 

BRITAIN: Scotland, Galloway, Laurieston, S.Snogerup & B.Snogerup 2272 (LD). NORWAY: Lofoten, 

C.Garcia-Herran 95-99 (NEU). 11. GERMANY: Baden, between Grötzingen and Weingarten, A.Kneucker 

[Cyper. Junc. Exs.] 57 (PR). POLAND: Babia Gora, 986 m, C.Garcia-Herran 92-781 (NEU; 2n=12). 

SLOVAKIA: Vysoké Tatry, Štrbské Pleso, 1370 m, V.Krajina [K.Domin & V.Krajina, Fl. Čechoslov. Exs.] 

335 (LD, PR, PRC). 12. FRANCE: Puy-de-Dôme, Giroux, Jalicon [Soc. Franc.] 7328 (PR). 13. BOSNIA: 

Glamoc, Mliniśte, Jul 1911, K.Vandas (PR). CROATIA: Plitvice Nat. Park, Sastavci, 500 m, N.Lundquist 

7729 (UPS). 34. TURKEY: Kastamonu, Ilgaz Dagh, 2080 m, P.H.Davis 21564 (BM). 

10b. Luzula sylvatica subsp. henriquesii (Degen) Pirajá, Agron. Lusit. 12: 359 (1951) 

Luzula henriquesii Degen, Magyar Bot. Lapok 5: 9 (1906); Luzula sylvatica var. henriquesii (Degen) Pau, Bol. 

Soc. Aragonesa Ci. Nat. 1916: 159 (1916). T: Portugal, Serra do Gerês, Ponte de Maceria, Moller 

[Fl. Lusit. Exs.] 850; holo: BP; iso: WU. 

33

34 


Figure 8. Luzula sylvatica subsp. sylvatica. A, habit; B, flower with unripe capsule; 

C, capsule; D, seed; E, tepal; F, stem T.S.; G, leaf T.S. (A–G, B.Błocki, Jul 1882, KRA). 

Scale bars: A = 5 cm; B–F = 2 mm; G = 1 mm. Drawn by T.Szynal. Reproduced with 

permission, from T.Szynal & J.Mądalski, Atlas Fl. Pol. 2/2: 153 (1931).


Luzula sylvatica var. paularensis Pau, Bol. Soc. Aragonesa Ci. Nat. 1916: 158 (1916). T: Spain, Madrid, 

Sierra de Guadarrama, El Paular, C.Vicioso; syn: MA 19634; 25 Aug 1912; C.Vicioso & Beltrán; syn: MA 

19641. 

Luzula sylvatica subsp. cantabrica P. Monts., Anales Inst. Bot. Cavanilles 21(2) [1963]: 441 (1964). 

T: Spain, Puerto Palombera, Reinosa, 1350 m, 26 Jun 1926, P.Font Quer; holo: BC. 

Plants usually (20–) 60–80 cm tall. Basal leaves (5–) 8–15 (–19) mm wide, gradually 

narrowed into long acute apex; margins ciliate. Inflorescence very lax; primary branches 

often elongated; secondary anthelas usually ±contracted, with 30–45 flowers; terminal 

clusters (1–) 2–3-flowered. Outer tepals usually (1.7–) 2.3–2.7 (–3.3) mm long. Anthers 

(0.7–) 0.9–1.8 (–2.1) mm long; filaments usually 0.3–0.6 mm long; style 1.9–2.2 mm long; 

stigmas 2.2–2.6 mm long. Capsule usually ±longer than perianth, ±ovoid, usually 1.6–2.6 mm 

long, mucro (0.2–) 0.4–0.7 mm long. Seeds usually 1.1–1.6 mm long, c. 0.6–0.7 mm wide; 

appendage c. 0.1–0.2 mm long. 2n=12, fide C.Garcia-Herran (see list of specimens); 

H.Nordenskiöld, Hereditas 37: 328–329 (1951). 

Confined to the northern part of the Iberian Peninsula. 12: POR, SPA. Found in woodlands, 

particularly along rivers, on damp soils. Map 28. 

12. PORTUGAL: Serra do Gerês, Albergaria, 700 m, C.Garcia-Herran 91-1134 (NEU; 2n=12); Serra da 

Estrela, Poço do Inferno, J.Matos, Cardoso & A.Marques 7905 (PR); Serra da Estrela: Lagoa do Covao do 

Curral, A.Fernandes, R.Fernandes & F.Sousa 4521 (LD); Alto Douro, Bragança, Pena de Águia, 750 m, 

P.Silva, B.Rainha & J.Martins 7788 (LD); Cinfaes, Fonte Nova, J.Paiva, J.Matos & A.Marques 8481 (PR). 

SPAIN: Asturias, Grado, San Pelayo, 2 May 1978, J.J.Lastra (FCO 24033); Asturias Luarca, along Esva R., 

near Pontigón, 24 Mar 1974, T.E.Díaz (FCO 23640); Salamanca, Miranda del Castañar, 690 m, 4 Apr 1985, 

D.Sánchez-Mata & V. de la Fuente (FCO 17362). 

A variable subspecies with several local variants. The eastern populations approach the type 

subspecies. 

10c. Luzula sylvatica subsp. sicula (Parl.) K.Richt., Pl. Eur. 1: 183 (1890) 

Luzula sicula Parl., Nuov. Gen. Sp. Monocot. 59 (1854); Luzula maxima subsp. sicula (Parl.) Nym., Consp. Fl. 

Eur. 751 (1882); Luzula sieberi subsp. sicula (Parl.) Pignatti, Fl. Ital. 3: 444 (1982), nom. inval. T: Sicily: 

‘monti di Sicilia nelle Madonie, in Montesoro, nei boschi di Caronia, di Mistretta, di Cannata, di Francavilla’; 

syn: n.v. 

Plants usually 30–45 (–65) cm tall. Basal leaves usually 3–4 (–6) mm wide, with elongated 

acuminate tip; margins and base very densely ciliate. Inflorescence lax, few-flowered; 

secondary anthela usually with (5–) 8–13 (–16) flowers; terminal clusters (1–) 2–3 (–4)flowered. 

Outer tepals usually 4.3–4.8 (–5.2) mm long. Anthers (1.7–) 2.2–2.5 (–3.0) mm 

long; filaments 0.2–0.6 mm long; style 1.7–1.9 mm long; stigmas on average 2.4–2.6 mm 

long. Capsule usually ±equalling perianth or shorter, ±ovoid, 3.5–4.1 (–4.4) mm long; mucro 

(0.2–) 0.4–0.8 mm long. Seeds (1.5–) 1.7–2.0 (–2.2) mm long, 1.1–1.3 mm wide; appendage 

c. 0.2 mm long. 2n=12, fide C.Garcia-Herran (see list of specimens). 

Southernmost Italy and Sicily. 13: ITA, SIC. In mountains, on volcanic slopes with sparse 

vegetation or broad-leaved (mostly Fagus sylvatica) woodlands. Map 29. 

13. ITALY: Calabria, E ridge of Montenero, 1400–1600 m, S.Snogerup & B.Snogerup 2081 (LD). SICILY: 

Messina, Floresta, 1350 m, Kraehembuhl 91-684 (NEU; 2n=12); Madonie, 600–1000 m, 18 Jul 1874, G.Strobl 

(K); Messina, Floresta, Mt Pojummoru, 1300 m, R.K.Brummitt, Hunt & Leistner 5104 (K); Valdemone, 

A.Todaro [Fl. Sic. Exs.] 649 (K, W); E and NE slopes of Mt Etna, 2250 m, 19–20 Aug 1873, G.Strobl (WU). 

10d. Luzula sylvatica subsp. sieberi (Tausch) K.Richt., Pl. Eur. 1: 183 (1890) 

Luzula sieberi Tausch, Flora 19: 423 (1836); Luzula maxima β [unranked] angustifolia Čelak., Prodr. Fl. 

Böhmen 1: 84 (1867), nom. illeg., non Roth (1788); Luzula sylvatica var. sieberi (Tausch) Buchenau, Bot. 

Jahrb. Syst. 12: 92 (1890); Luzula maxima subsp. sieberii (Tausch) P.Fourn., Fl. Compl. Plaine Franç. 527 

(1928). T: [Austrian Alps] F.W.Sieber [Herb. Fl. Austr.] 104, ut Juncus maximus β angustifolius; lecto: 

PRC, designated here; isolecto: BRNM. 

Illustration: S.Pignatti, Fl. Ital. 3: 444 (1982). 

35

36 


Plants usually up to 55 (–70) cm tall. Basal leaves usually 4–8 (–9) mm wide; apex 

subobtuse, sharply mucronate; margins ciliate. Inflorescence lax, ±few-flowered; secondary 

anthelas usually with 15–20 flowers; terminal clusters 1–3-flowered. Outer tepals usually 

2.7–3.9 mm long. Anthers usually 1.7–1.9 (–2.3) mm long; filaments 0.2–0.5 mm long; style 

c. 1.9–2.2 mm long; stigmas 2.2–2.6 mm long. Capsule usually ±equalling or shorter than 

perianth, oblong-ovoid, usually (2.3–) 2.8–3.0 (–3.3) mm long; mucro (0.2–) 0.7–0.8 mm 

long. Seeds 1.2–1.9 mm long, c. 0.7 mm wide; appendage c. 0.2 mm long. 2n=12, fide 

Druskovic, Int. Organ. Pl. Biosyst. Newslett. 24: 14 (1995). 

South-Central Europe, most frequently in the Alps and N Italy. 11: AUT, GER, SWI. 

12: FRA, SPA. 13: ITA, YUC. Found in mountain coniferous woodlands, in tall-grass 

vegetation near timberline. Map 30. 

11. AUSTRIA: Karnische Alpen, Kleiner Pal, c. 1800 m, L.Kirschnerová 4-989 (PRA); Tyrol, Luttach, 

Schönberg, 1 Jul 1896, Treffer (PR); Tyrol, Brenner, 1500–1600 m, R.Huter (PR, WU). SWITZERLAND: 

Graubünden, Versam, Maiensäss Fahn, 1200 m, L.Meisser [J.Braun-Blanquet, Fl. Raet. Exs.] 421 (PR). 

13. ITALY: Etruria, Grosseto, Mt Amiata, 800–1200 m, A.Fiori & A.Béguinot [Fl. Ital. Exs.] 2629 (WU). 

Occasional intermediates exist in regions of contact with subsp. sylvatica and in marginal 

parts of the range, (for instance in the E Pyrenees). 

Presumed Hybrids 

Luzula ×hermannii-muelleri Asch. & Graebn., Syn. Mitteleur. Fl. 2(2): 507 (1904) = Luzula 

luzuloides (Lam.) Dandy & E.Willm. × L. sylvatica (Huds.) Gaudin 

T: [Germany] Westfalen, Arnsberger Wald südlich von Drüggelte, H.Mueller; syn: n.v. 

A rather doubtful hybrid requiring further study. 

Luzula ×favratii K.Richt., Pl. Eur. 1: 185 (1890) = Luzula luzuloides (Lam.) Dandy & 

E.Willm × L. nivea (Nathh.) DC. 

T: [Switzerland] Bois de Sauvabelin, Lausanne, L.Favrat; syn: W. 

Luzula × schultzorum Dalla Torre & Sarnth., Farn- u. Blütenpfl. Tirol. 1: 434 (1906). 

T: [Austria] 'B, Ratzes'; syn: n.v. 

A rare hybrid ±intermediate between the parents; known from Austria and Switzerland. 

It requires further study. 

Luzula ×levieri Asch. & Graebn., Syn. Mitteleur. Fl. 2(2): 509 (1904) = L. nivea (Nathh.) DC. 

× Luzula pedemontana Boiss. & Reut. 

T: [Italy] in den Etrurischen Apenninen, im Walde Della Fornace bei Boscolungo, E.Levier; syn: n.v. 

A rather doubtful hybrid requiring a field investigation. 

Luzula ×lepetitiana Beyer, Verh. Bot. Vereins Prov. Brandeburg 41: xiv (1900) = Luzula 

pedemontana Boiss. & Reut. × L. sylvatica (Huds.) Gaudin 

T: [N Italy] Borgone, Sangone, am nordöstlichen Abhange des Mte. Salancia, R.Beyer; holo: n.v. 

Luzula ×hasleri Murr, Magyar Bot. Lapok 28: 68 (1930) = Luzula alpinopilosa (Chaix) 

Breistr. × L. sylvatica subsp. sieberi (Tausch) K.Richt. 

T: Liechtenstein, Heupiel (Heubühl), 1867 m, 2 Aug 1921, J.Murr; syn: n.v. 

Material not seen. The existence of the hybrid is rather doubtful.


Luzula ×johannis-principis Murr, Liechtenst. Volksblatt, 1921, no 49 [21 Jun 1921] = Luzula 

sylvatica subsp. sieberi (Tausch) K.Richt. × L. alpina Hoppe 

T: Liechtenstein, Rojaberg, 1630 m, J.Murr; syn: K, S, W. 

Numerous specimens from the type locality represent ±sterile plants, partly referable to 

L. multiflora, partly of an appearance intermediate between the presumed parents. Further 

study is needed. 

Luzula ×pfaffii Murr, Allg. Bot. Z. Syst. 1910: 118 (1910) = Luzula lutea (All.) DC. × 

L. luzuloides subsp. rubella (Mert. & W.D.J.Koch) Holub 

T: [Italy] Bozen, Ritterhorn, 2260 m, 4 Sep 1909, W.Pfaff; syn: n.v. 

A rather doubtful hybrid; further study is needed. 

Luzula ×bornmuelleriana Kük., Mitth. Thüring. Bot. Vereins 23: 90 (1908) = Luzula 

alpinopilosa × L. lutea 

T: [Austria] Tirol, Pitztal, Mittelberg, Taschachbach, 2000 m, G.Kükenthal; syn: n.v. 

Another very improbable hybrid, Luzula pilosa (L.) Willd. × L. sylvatica (Huds.) Gaudin, has 

been reported under an invalid name, L. × buchenaui P.Fourn., Quatre Fl. France 154 (1935). 

Sect. 2. ATLANTICAE 

Luzula sect. Atlanticae Kirschner, Taxon 50: 1107 (2001) 

T: Luzula atlantica Braun.-Blanq. 

Perennials with short ascending rhizomes. Leaf tip acute. Lower bract subherbaceous, usually 

±membranous, shorter than inflorescence. Inflorescence racemose, decompound, with 3–8 

thin primary branches; secondary branches rare; flower heads subglobose, (1–) 2–7-flowered. 

Tepals membranous, mucronate, with a castaneous midrib. Style very short. Seeds subglobular, 

smooth; appendage indistinct. 2n=12. 

A monotypic section restricted to the mountains of Morocco. 

11. Luzula atlantica Braun-Blanq., Vierteljahrsschr. Naturf. Ges. Zürich 73, Suppl. 15: 

347 (1928) 

T: Morocco, Great Atlas, Zerekten, 1500–2000 m, J.Braun-Blanquet; holo: MPU, n.v. [In the Braun-Blanquet 

herbarium, MPU, no specimen with the above locality was located. The plant photographed in the protologue, 

deposited in MPU and labeled 'Gourdouze, 2000 m' thus may be a good candidate for a lectotype.] 

Illustrations: J.Braun-Blanquet, Vierteljahrsschr. Naturf. Ges. Zürich 73, Suppl. 15: fig. 7, 356 (1928); 

R.Maire & Weiller, Fl. Afr. Nord 4: 308, fig. 729 (1957). 


Perennials 10–30 (–35) cm tall, caespitose, ciliate; rhizome short, vertical to oblique, sometimes 

many-headed; stolons absent. Basal leaves usually 4–10 (–14) cm long, 2–4 (–5) mm 

wide; cauline leaves 1 (–2), usually 2–5 cm long. All leaves with long acuminate tip; 

margins papillose-serrulate. Lower bract ±herbaceous, sometimes ±membranous, shorter than 

inflorescence. Inflorescence decompound; main branches 3–8, smooth or sparsely papillose 

below clusters, thin, c. 0.2 mm in diam., often flexuose, patent to erect; secondary branches 

rare, with each branch bearing loose terminal (1–) 2–7-flowered glomeruliform clusters 4–5 mm 

in diam. Bracteoles ovate-lanceolate, subaristate, glabrous to sparsely ciliate, 1.5–2.0 mm 

long, membranous-scarious; midrib often castaneous-brown. Tepals ±equal, glabrous, entire 

or the inner ones minutely denticulate below apex, lanceolate to ovate-lanceolate, often 

almost entirely membranous with dark castaneous-brown midrib and short mucro, sometimes 

castaneous-brown with broad membranous margins, 1.9–2.6 mm long. Stamens 6; anthers 

c. 0.4 mm long, much shorter than filaments; filaments 0.8–0.9 mm long; style c. 0.2 mm 

37

38 


long; stigmas 1.0–1.2 mm long. Capsule subglobular, ±obtuse, mucronate, castaneous-brown; 

capsule segments 1.7–2.0 mm long, including mucro 0.1–0.2 mm long and 1.4–1.5 mm wide. 

Seeds subglobular, pale castaneous-brown, smooth, shining, 0.8–0.9 (–1.0) mm long, 0.7–0.8 mm 

wide; apex pale; basal appendage indistinct, to 0.1 mm long, very sparsely fibrillate. 2n=12, 

fide N.Galland, Trav. Inst. Sci. Univ. Mohammed V, Sér. Bot., 35 (22): 106–121 (1988). 

Endemic to the Great Atlas and Anti-Atlas in Morocco. 20: MOR. Amongst shady mountain 

rocks in open oak woodlands. Map 31. 

20. MOROCCO: Great Atlas, Oukaimeden, 2500–2600 m, P.H.Davis 67595 (E); Reraya, Sidi-Chamarouch, 

2350–2500 m, 1923, R.V.Litardiére (MPU); Great Atlas, Reraya Valley, Sidi-Chamarouch, 2400–2500 m, 

21 Jul 1924, R.Maire (P); Great Atlas, Gourdouze, 1800 m, Apr 1928, J.Braun-Blanquet & I.Braun-Blanquet 

(MPU). 

A species in many respects intermediate between sections Anthelaea and Diprophyllatae. 

The seed characters clearly point to Luzula (Marlenia) elegans Lowe. 

Sect. 3. NODULOSAE 

Luzula sect. Nodulosae Chrtek & Křísa, Preslia 46: 211 (1974) 

Luzula subsect. Nodulosae (Chrtek & Křísa) Novikov, Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 95(6): 

66 (1990). T: Luzula nodulosa E.Mey. 

Perennials with thick nodulose horizontal rhizomes. Leaf tips acuminate, subulate. Lower 

bract ±membranous to subherbaceous. Inflorescence umbelloid or anthelate, composed of 

capituliform dense large clusters, on ±suberect peduncles. Seeds broadly ovoid, large, 

1.7–1.9 × 1.3–1.4 mm, with apical and basal patches of looser outer seed-coat; appendages 

absent; base distinctly fibrillate. 

A monotypic section confined to the Mediterranean region, disjunctly in NW Africa and 

from Greece to W Anatolia. 

12. Luzula nodulosa [Bory & Chaub.] E.Mey., Linnaea 22: 410 (1849), nom. cons. 

propos. 

Juncus nodulosus Bory & Chaub., Exp. Sci. Morée Bot. 3: 105, t. 11, f. 1 (1832), nom. illeg., non Wahlenb.; 

Juncus graecus Chaub. & Bory, Nouv. Fl. Pélop. 23, Plate 12, fig. 1 (1838); Luzula graeca (Chaub. & Bory) 

Kunth, Enum. Pl. 3: 310 (1841); Juncoides nodulosa (E.Mey.) Kuntze, Revis. Gen. Pl. 2: 724 (1891); Luzula 

nodulosa var. graeca (Chaub. & Bory) R.Maire, Bull. Soc. Hist. Nat. Afrique N. 22: 319 (1931). T: [Greece, 

Pylos, Arcadia] L.A.Chaubard; holo: P. 

Luzula nodulosa var. mauretanica R.Maire & Trab., in R.C.J.E.Maire, Bull. Soc. Hist. Nat. Afrique N. 22: 319 

(1931). T: Algeria, Monts Tlemcen, entre Terni & Sebdou, L.Trabut; holo: P. 

Illustration: F.G.P.Buchenau, in H.G.A.Engler, Pflanzenr. (iv.36) 25: 65, fig. 46a, 66, fig. 46b (1906). 

Perennials c. (15–) 30–55 (–70) cm tall, loosely caespitose or one-stemmed; rhizome thick, 

c. 3–5 mm in diam., creeping to ascending, often with fibrous cover, nodulose, with internodes 

usually 0.5–2.0 cm long; stolons short, ascending. Lowermost leaves cataphyllous, c. 1–3 cm 

long, brownish; basal leaves flat, 7–16 (–20) cm long, 3.5–5.0 (–7.5) mm wide; cauline 

leaves 3–4 (–5), (1.5–) 2.5–5.0 (–15) cm long; margins densely ciliate, later sometimes 

glabrescent, densely papillose-serrulate; apex subulate. Lower bract brownish-membranous 

to subherbaceous, c. 1.0–3.5 (–5.0) cm long, shorter than inflorescence. Inflorescence of 

(1–) 2–5 (–15) large hemispherical to cup-shapedclusters; clusters (2–) 3–5 (–11)-flowered, 

0.7–1.0 cm long, c. 1.0–1.5 cm wide; peduncles suberect, c. 2–10 cm long, smooth; 

secondary branches absent or frequent, distally inserted, short. Bracteoles scarious, often 

brownish at base, ovate, acuminate, c. 3.0–3.5 mm long; midrib distinct. Tepals ±equal, 

ovate-lanceolate or broadly lanceolate, c. 5.0–6.5 mm long, acute to acuminate; central band 

deep castaneous-brown; margins distinct, broad, membranous. Stamens 6; anthers 1.7–3.5 mm 

long, 2–10 times as long as filaments; filaments 0.3–0.9 mm long; style 1.8–2.7 mm long; 

stigmas c. 2.5–4.0 mm long. Capsule ovoid to broadly ovoid, acuminate, shorter than 

perianth, dark castaneous to pale castaneous-brown; capsule segments 3.3–4.0 × 2.3–2.8 mm


long; mucro to 0.5 mm long. Seeds broadly ovoid, deep brown, 1.7–1.9 mm long, 1.3–1.4 mm 

wide, with apical and basal whitish patches of loose outer seed-coat, distinctly fibrillate. 

Disjunctly in two areas: the E Mediterranean (from Greece to W Anatolia) and NW Africa 

(Algeria and Morocco). 13: GRC, KRI. 20: ALG, MOR. 34: EAI, TUR. Rocky slopes, open 

woodlands. Map 32. 

13. GREECE: Attica, 650–1300 m, T.H.Heldreich [Herb. Graecum Norm.] 148 (P); Attica, Mt Pentelica, 

T.G.Orphanides 1105 (WU). KRITI: Askoyplicus, 1846, T.H.Heldreich (W); Hag. Vasilis, Kedros, 

K.H.Rechinger 849 (WU); Karpathos, Mt Kolla, W.Greuter 5314 (W). 20. ALGERIA: Kabylie, Mt Magris, 

1700 m, E.Reverchon [Pl. Alger.] 364 (P, WU); Djurdjura, Ras Timedouine, 6 km E of Tikdja, 1860–1900 m, 

P.H.Davis 59408 (E). MOROCCO: Bab-Bou-Idir, C.G. van Steenis 19181 (L). 34. EAST AEGEAN IS.: 

Samos, Mt Ambelos, 800 m, K.H.Rechinger & F.Rechinger 3905 (K). TURKEY: Çanakkale, Erenköy, 

Renkoei Scala, 16 Apr 1883, P.E.E.Sintenis [It. Trojan.] 297 (W, UPS). 

Variable in stamen and pistil characters. There are two main forms, one with anthers 

1.7–2.5 mm long, about 2–3 times as long as filaments, and style c. 2 mm long, the other with 

anthers 2.0–3.5 mm long, usually 4–8 times as long as filaments, and style c. 2.2–2.7 mm 

long. In N Africa, and in Morocco in particular, plants with longer anthers and styles 

predominate, mostly on limestone. Many African plants, however, are indistinguishable from 

the E Mediterranean ones, and intermediates are not rare. 

Sect. 4. DIPROPHYLLATAE 

Luzula sect. Diprophyllatae Satake, in T.Nakai & M.Honda, Nov. Fl. Jap. 1: 25 (1938) 

T: Luzula wahlenbergii Rupr.; lecto, designated here. 

Luzula ser. Effusae Satake, in T.Nakai & M.Honda, Nov. Fl. Jap. 1: 25 (1938); Luzula sect. Effusae [Satake] 

K.F.Wu, J. E. China Norm. Univ., Nat. Sci. Ed., 1992(3): 91 (1992). T: Luzula effusa Buchenau. 

Luzula ser. Spadiceae Satake in T.Nakai & M.Honda, Nov. Fl. Jap. 1: 26 (1938). T: Luzula wahlenbergii Rupr. 

Luzula sect. Glabratae Chrtek & Křísa, Preslia 46: 211 (1974). T: Luzula glabrata (Hoppe) Desv. 

Luzula subsect. Parviflorae Kovt., Bot. Žurn. (Moscow & Leningrad) 72: 1398 (1987); Luzula sect. 

Parviflorae [Kovt.] K.F.Wu, J. E. China Norm. Univ., Nat. Sci. Ed., 1992(3): 92 (1992). T: Luzula 

parviflora (Ehrh.) Desv. 

Perennials, subglabrous to sparsely ciliate; rhizome branched, ascending. Leaves with acute 

tips. Lower bract herbaceous to subherbaceous. Inflorescence decompound, a ±nodding to 

suberect panicle. Flowers usually borne singly, some in 2–5-flowered clusters. Tepals 

usually 2.0–2.5 (–3.5) mm long. Seeds usually ellipsoid to oblong-ellipsoid; appendage 

absent or, when visible, basal, fibrillate. 

A section with 12 species of mostly subarctic and oro-boreal, or alpine distribution. Main 

centres of diversity are in W North America and Europe; two species reach the southern 

hemisphere. Almost exclusively, diploids (2n=12 AL) and agmatoploids with 2n=24 BL are 

known in the section. The species are very close to one another in their general habit. 

1 Upper part of capsule and mucro densely papillose; inflorescence usually 

storied (inflorescence subdivided in several subequal panicles separated 

by long rhachis segments; inflorescence usually constitutes 1/3 of plant 

height or more) 

23. L. effusa 

1: Capsule smooth; inflorescence storied or not so 

2 Inflorescence storied 13. L. gigantea 

2: Inflorescence not storied 

3 Panicle branches abbreviated, and inflorescence therefore 

subcongested 16. L. subcapitata 

3: Panicle branches (at least the main ones) long; panicle loose or 

subdense, never subcongested 

39

40 


4 Tepals stramineous to pale brownish, with apex often ±reflexed; 

inflorescence very lax, with rigidly divaricate branches 17. L. divaricata 

4: Tepals blackish brown to dark castaneous-brown, with apex not 

reflexed; inflorescence lax to subdense, ±nodding to suberect, with 

branches not rigidly divaricate 

5 Basal leaves densely to sparsely ciliate. [Mexico to Panama] 14. L. denticulata 

5: Basal leaves subglabrous. [Outside C America] 

6 Capsule segments usually 3.0–3.5 × 1.8–2.0 mm 19. L. hitchcockii 

6: Capsule segments to 2.9 mm long and to 1.6 mm wide 

7 Anthers ±equalling filaments or shorter; style up to 0.5 mm long 

8 Plants usually more than 30 cm tall; cauline leaves (3–) 4–5, 

usually 5–10 mm wide 22. L. parviflora 

8: Plants usually up to 30 cm tall; cauline leaves 1–3, usually 

less than 5 mm wide 

9 Cauline leaves bluish green to greyish green, (2–) 3; lower 

bract 1.5–2.5 cm long 18. L. piperi 

9: Cauline leaves mid-green, 1–2; lower bract to 1 cm long 15. L. wahlenbergii 

7: Anthers (2–) 3–5 (–9) times longer than filaments; style usually 

longer than 0.5 mm 

10 Capsule segments to 2.4 mm long; seeds stramineous to pale 

brown 

10: Capsule segments 2.4–3.0 mm long; seeds brown to dark 

brown 

11 Cauline leaves 10–20 cm long, usually 3–6 mm wide; anthers 

24. L. alpinopilosa 

usually 3–4 times longer than filaments 

11: Cauline leaves up to 6 cm long, usually 4.5–8.0 mm wide; 

21. L. desvauxii 

anthers usually 5–7 times longer than filaments 20. L. glabrata 

13. Luzula gigantea Desv., J. Bot. (Desvaux) 1: 145 (1808) 

Juncoides spadicea Kuntze var. gigantea (Desv.) Kuntze, Revis. Gen. Pl. 2: 724 (1891); Juncoides gigantea 

(Desv.) E.Sheld., Minnesota Bot. Stud. 1: 62 (1894). T: Colombia, in montibus humidis Quindio in regno 

Santa-fe de Bogota, A.Humboldt & A.J.Bonpland; holo: P. 

Luzula paniculata Desv., J. Bot. (Desvaux) 1: 147 (1808). T: ‘In America calidiore’; holo: P. 

Illustrations: M.Barros, Darwiniana 10: 318, fig. 10 (1953); H.Balslev, Fl. Neotrop. Monogr. 68: 53, fig. 26 

(1996). 

Map: H.Balslev, Fl. Neotrop. Monogr. 68: 54, fig. 27 (1996). 

Perennials 20–90 cm tall; rhizome ascending, 1.5–3.0 mm in diam.; stolons long, ascending. 

Basal leaves 10–30 cm long, (5–) 10–20 mm wide, flat; margin densely papillose-serrulate, 

ciliate in young leaves; cauline leaves usually 2–4, with upper ones usually up to 10–13 mm 

wide; sheaths to 6 cm long; leaf tip acute. Inflorescence a rich loose compound panicle 

constituting more than one third of plant height, storied and subdivided by 5–15 cm long 

rhachis segments into 4–5 subequal nodding anthelate parts; flowers borne singly (rarely in 

clusters of 2–3). Bracts of primary inflorescence branches resembling cauline leaves, with 

proximal ones to 20 cm long and distal ones gradually shorter; bracts on distal branches 

brownish-membranous, linear-lanceolate, c. 1.0–1.5 cm long. Bracteoles ovate to ovatelanceolate, 

brownish-membranous, 1.6–2.0 mm long, usually entire, glabrous or sparsely 

ciliate, inserted 0.2–0.5 mm below flowers. Tepals ±unequal, glabrous, acuminate, usually 

dark castaneous brown, margins paler; outer tepals usually 2.2–2.7 mm long; inner tepals 

usually 2.0–2.5 mm long. Stamens 6, 1.0–1.5 mm long; anthers 0.3–0.6 mm long; filaments 

0.7–0.9 mm long; style 0.2–0.3 mm long; stigmas ±patent, 0.6–0.8 mm long. Capsule broadly 

ellipsoidal, apiculate, shorter than perianth, castaneous, shining; capsule segments often slightly


mucronate, 1.6–1.9 × c. 1.1 mm long. Seeds ±ellipsoidal, (0.9–) 1.0–1.1 × 0.5–0.6 mm; outer 

seed-coat white to light brown; appendage almost not visible, c. 0.1 mm long; base fibrillate. 

Fig. 9. 

South America, in the Andes from Venezuela and Colombia to Bolivia and N Argentina. 

82: VEN. 83: BOL, CLM, ECU, PER. 85: AGW. Open mountain slopes, páramo, along 

streams, usually between 3000 and 4000 m. Map 33. 

82. VENEZUELA: Mérida, Trujillo, J.Linden 412 (BM, BR, G, P, US). 83. BOLIVIA: Cochabamba, 

Chapare road at km 110, 3000 m, R.F.Steinbach 611 (LPB, NY, S, US, WIS); La Paz, Larecaja, near Sorata 

and Mt Sanquiarie, 3200 m, G.Mandon 1423 (BM, K, NY, P, S, US). COLOMBIA: Cauca, Cordillera 

Central, Puracé, 3450 m, K.Sneidern 1740 (S); Boyaca, Sierra Nevada de Cocuy, Quebrada de las Playas, 4100 

m, P.J.Grubb et al. 321 (K, US). ECUADOR: Imbabura, Lago San Marcos Cayambe, Wash et al. 5460 (K); 

Imbabura, Mojanda, 10 km SSW Otavalo, 2900–3150 m, B.Sparre 13476 (S); Pichincha, Mt Corazón, 

E.Asplund 9684 (S). PERU: Huacás, Santiago de Chuco, 2800 m, A.Sagástegui, J.Mostacero & Diestra 

11913 (AAU, MO, K, NY, F); Mito, c. 3000 m, J.F.Macbride 3367 (S). 

At first sight very similar to SE Asian Luzula effusa (the only other taxon having storied 

inflorescence) but differs from it in having smooth capsules, longer outer tepals, anthers 

substantially shorter than filaments, narrower seeds and longer bracteoles. 

14. Luzula denticulata Liebm., Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 

2: 46 (1850) 

Luzula parviflora var. denticulata (Liebm.) Buchenau, Bot. Jahrb. Syst. 7: 171 (1885). T: Mexico, Orizaba, 

Vaqueria del Jacal, 3050 m, F.Liebmann; lecto: C, fide L.Hämet-Ahti, Ann. Bot. Fenn. 8: 370 (1971); isolecto: 

C, G, HAL, K, P, S, UPS. 

Luzula laetevirens Liebm., Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 2: 46 (1850). T: Mexico, 

Orizaba, 2750–3050 m, F.Liebmann; lecto: C, fide L.Hämet-Ahti, Ann. Bot. Fenn. 8: 370 (1971); isolecto: C, P. 

Luzula latifolia Liebm., Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 2: 47 (1850). T: Mexico, 

Oaxaca, Cumbre de Ixtepec, 3050 m, F.Liebmann; lecto: C, fide L.Hämet-Ahti, Ann. Bot. Fenn. 8: 370 (1971); 

isolecto: C. 

Luzula gigantea var. vulcanica Woodson, in R.E.Woodson & R.J.Seibert, Ann. Missouri Bot. Gard. 26: 278 

(1939). T: Panama, Chiriquí, Loma Larga, Volcán Chiriquí, 2500–3380 m, R.E.Woodson jr., P.H.Allen & 

R.J.Seibert 1094; holo: MO; iso: GH, NY, US, photo AAU. 

Illustration: H.Balslev, Fl. Neotrop. Monogr. 68: 56, fig. 28 (1996). 


Perennials 15–80 cm tall; rhizome ascending, long, often with long ascending stolons. Basal 

leaves 5–25 cm long, 8–12 mm wide; cauline leaves usually 3–5 (–8), usually 6–8 cm long, 

4–5 mm wide. All leaves usually densely ciliate, less often subglabrous, acute, with ±papilloseserrulate 

margins. Lower bract herbaceous, up to 6 cm long; next upper bract linear, brownish 

membranous, ciliate, c. 1 cm long. Inflorescence terminal, a loose nodding anthelately 

arranged panicle 5–20 × 3–10 cm; flowers borne singly. Bracteoles brownish-membranous, 

±ovate, acute, 1.2–1.6 mm long, ciliate, inserted c. 0.1–0.2 mm below flower. Tepals 

subequal, glabrous, narrowly-lanceolate, acuminate, pale to dark brown, 1.7–2.4 mm long. 

Stamens 6; anthers 0.4–0.7 mm long, equalling or longer than filaments; filaments 0.3–0.5 mm 

long; style 0.1–0.2 mm long; stigmas 0.8–1.0 mm long. Capsule dark brown, shining, ovoid, 

acute, often equalling or exceeding perianth; capsule segments 1.5–1.7 × 1.0–1.3 mm. Seeds 

oblong-ellipsoidal, brown, 1.0–1.1 mm long, 0.6–0.7 mm wide; appendage indistinct, to 

c. 0.1 mm, fibrillate. Fig. 10. 

Confined to Central America, from C Mexico to Panama. 79: MXC, MXE, MXG, MXS, MXT. 

80: COS, GUA, PAN. Grasslands, pastures and open woodlands, between 200–3900 m. 

Map 34. 

79. MEXICO CENTRAL: Distrito Federal, Contreras, Los Dinamos, 2800 m, A.Ventura 3580 (AAU); Distrito 

Federal, Mt Ajusco, 3840 m, J.H.Beaman 2806 (DS, MICH, US); México State, La Ciénega, Iztaccíhuatl, 

R.Galván 735 (AAU); Puebla, Sierra Negra, Mt Orizaba, 3200 m, E.K.Balls et al. 4475 (K, UC, US). 

MEXICO NORTHEAST: Hidalgo, Epazoyucan, Nopalillo, 2850 m, R.Galván 580 (AAU); Hidalgo, El Chico, 

E.Lyonnet 265 (CAS, GH, MEXU, MO, NY, K, US). 80. COSTA RICA: Cartago, Volcán Irazú, 3000 m, 

K.Barringer, Christenson & L.D.Gomez 3273 (AAU); Cartago, Cerro de la Muerte, 3000–3500 m, 

41

42 


Figure 9. Luzula gigantea. Note wide leaves, storied inflorescence, single inserted flowers. 

(Fagerlind & Wibom 1405 bis , S). Scale bar: 5 cm. Drawn by K.Tind. Originally published in 

G.Harling & B.Sparre (eds.), Fl. Ecuador 11: 10, fig. 2 (1979), reproduced with permission, 

from H.Balslev, Fl. Neotrop. Monogr. 68: 53, fig. 26 (1996), © Lunds Botaniska Förening, 

© 1996, The New York Botanical Garden.


Figure 10. Luzula denticulata. A, habit; B, flowers with fruits. (A, B, Nelson 645, US). 

Scale bars: A = 5 cm; B = 2 mm. Drawn by K.Tind. Reproduced with permission, from 

H.Balslev, Fl. Neotrop. Monogr. 68: 56, fig. 28 (1996), © 1996, The New York Botanical 

Garden. 

43

44 


R.W.Holm & H.H.Iltis 485 (A, MO, P); Irazú, La Playita, 3300 m, Pettier 14125 (AAU). GUATEMALA: 

Huehuetenango, San Mateo Ixtatán, Sierra de los Cuchumatanes, 2900 m, D.E.Breedlove 11555 (DS, LL, 

MICH, US). 

In Panama and S Mexico, plants intermediate between L. denticulata and L. gigantea occur, 

but are rare. They are almost glabrous and have ±storied inflorescence (vouchers in AAU). 

Further study is necessary. 

15. Luzula wahlenbergii Rupr., Fl. Samojed. Cisural. 58 (1845) 

T: North Russia, ‘Litt. austr. ins. Kolguev’, F.J.Ruprecht; syn: LE, UPS, W; Norway, Finnmark, Kaafjord, 

M.Blytt in E.Fries [Herb. Norm. fasc. VIII] 70; syn: UPS, S. 

Luzula spadicea var. kunthii E.Mey., Linnaea 22: 403 (1849). T: Sweden, Lapponia Pitensis, 

G.Wahlenberg; syn: W;. Norway, Finnmark, Kaafjord, M.Blytt in E.Fries [Herb. Norm. fasc. VIII] 70; syn: 

UPS, S. 

Luzula borealis Fr., Summa Veg. Scand. 219 (1846). T: ‘Per alpes Scandinaviae altioris totius’. Norway, 

Finnmark, Kaafjord, M.Blytt in E.Fries [Herb. Norm. VIII] 70; syn: UPS, S. 

Illustration: J. Lid, Norsk & Svensk Fl. 202, fig. 99a (1963). 

Maps: L.Hämet-Ahti, Aquilo, Ser. Bot., 3: 18, fig. 5 (1965); E.Hultén & M.Fries, Atlas N. Eur. Vasc. Pl. 

1: map 167 (1986). 

Perennials c. 20–30 cm tall, subglabrous, caespitose, sometimes with short ascending 

vegetative shoots. Stem with several basal and 1–2 cauline flat leaves; basal leaves 5–10 cm 

× 3–5 mm; upper leaves up to 3–5 cm × 2–4 mm; leaf margins papillose-serrulate; leaf tip 

acute. Inflorescence a lax nodding ‘panicle’ of 10–30 flowers, c. 4 cm long and 3 cm wide; 

flowers borne singly, pedunculate, or in 2(–3)-flowered clusters. Lower bract 4–10 mm long. 

Bracteoles up to 2.0 mm long, fimbriate-ciliate near apex. Tepals equal, 2.0–2.8 mm long, 

±acuminate, entire, ±brown. Stamens 6; anthers 0.4–0.6 mm long, ±equalling filaments; style 

up to 0.5 mm long; stigmas c. 1 mm long. Capsule ellipsoidal, ±subacuminate, ±exceeding 

tepals; capsule segments brown, 2.0–2.4 × 1.1–1.5 mm. Seeds narrowly ellipsoidal, 1.2–1.4 × 

c. 0.7 mm, dark castaneous brown, without appendage; base fibrillate. 2n=24 [BL], 


Arctic Eurasia and mountains of Scandinavia, Russian Far East and Transbaicalia, Alaska 

and Arctic Canada and Greenland. 10: FIN, NOR, SVA, SWE. 14: RUN. 30: BRY, CTA, 

KRA, WSB, YAK. 31: KAM, KHA, KUR, MAG, PRM. 70: ASK, GNL, NWT, YUK. 

72: QUE. Arctic and alpine mossy tundra, wet grasslands. Map 35. 

10. FINLAND: Lapponia enontekiensis, 69°N, Aug 1912, J.Montell s.n. (PR). NORWAY: Finnmark, Nord- 

Varanger, Fossefjellet, N.Lundquist 4944 (UPS). SWEDEN: Torne Lappmark, Jukkasjärvi, 24 Jul 1904, 

Ringselle s.n. (PR). 14. RUSSIA NORTH: Murmansk, Imandra Lake, R.R.Pohle 6980 (MW). 30. WEST 

SIBERIA: Tyumen, Distr. Priuralskiy, 2.5 km N of Shuchye. 31. RUSSIAN FAR EAST: Magadan Distr., 

Pitlekaj, 1879 F.Kjellman s.n. (S); Chukotka, Nedle, A.A.Krause 175 (UPS). Kamchatka, Pal’matkina R., 

1932 B.N.Gorodkov & B.A.Tikhomirov s.n. (S). 70. ALASKA: Nunivak Is., Nash Harbour, 1938 

J.P.Anderson 3878, 3875 (S); Brooks Range, De Long Mtns, Feniak Lake, 600 m, K.Holmen & O.Mårtensson 

61-1130 (UPS). CANADA, NORTHWEST TERRITORIES [Nunavut]: Franklin Distr., Frobisher Bay, 1959, 

E.Hultén s.n. (S). 

Close to L. parviflora and L. piperi. From the former it can be distinguished by lower 

growth, narrower leaves, 1–2 cauline leaves; from the latter by linear-lanceolate cauline 

leaves and dark brown seeds (L.Hämet-Ahti, Aquilo, Ser. Bot., 3: 11–21, 1965). 

16. Luzula subcapitata (Rydb.) H.D.Harr., Man. Pl. Colorado 151 (1954) 

Juncoides subcapitata Rydb., Bull. Torrey Bot. Club 1904: 401 (1904). T: USA, Colorado, Silver Plume, 

1895, C.L.Shear 4611; holo: NY. 

Illustration: J.Coffey Swab, Fl. North Amer. 22: 262 (2000). 

Map: J.Coffey Swab, Fl. North Amer. 22: 261 (2000). 

Perennials 15–30 cm tall, loosely caespitose; rhizome ascending, with numerous short 

ascending vegetative shoots. Stem ±green. Basal leaves 1–4, 3.5–6.0 cm long, 4–5 mm wide; 

cauline leaves 3 (–4), with lower ones broader, 3–6 cm long, 2.0–5.5 mm wide. All leaves


apiculate, glabrous or with sparse hairs at sheath mouth. Lower bract up to 1.5 cm long, 

rarely 3 cm long, sometimes overtopping inflorescence. Inflorescence a single congested 

dark head composed of indistinct clusters; each flower usually with a short visible peduncle to 

1 mm. Bracteoles 1.0 (–1.5) mm long, brown; apex obtuse, ciliate. Tepals equal, glabrous, 

ovate-lanceolate, 1.8–2.2 mm long, ±dark castaneous-brown; margins not visible. Stamens 6; 

anthers (0.3–) 0.4 mm long, slightly shorter than or equal to filaments; filaments c. 0.4 mm 

long; style 0.2–0.3 mm long; stigmas 0.8–1.0 mm long. Capsule subglobular-ovoid, equalling 

or slightly exceeding perianth, almost rounded above, with a short apiculate mucro; capsule 

segments 1.7–1.8 × 1.2–1.3 mm long. Seeds paler castaneous-brown, c. 1.0 mm long, 

c. 0.6 mm wide; appendage absent; base fibrillate. 

A restricted endemic of mountains in Colorado, USA. 73: COL. Damp, boggy areas near 

alpine lakes and streams, usually above 3000 m. Map 36. 

73. USA, COLORADO: San Juan Co., Tenmile Basin, Animas R. drainage, SE of Silverton, Needle Mtn 

Quadrangle, J.Michener 789 (S); Willow Lake, W.W.Robbins 7933 (H). 

Luzula subcapitata occupies an isolated position in the section; it is the only species with 

abbreviated inflorescence branches and peduncles, i.e., with a ±congested inflorescence. 

17. Luzula divaricata S.Watson, Proc. Amer. Acad. Arts (n.s.) 6: 302 (1879) 

Juncoides divaricata (S.Watson) Coville, Contr. U. S. Natl. Herb. 4: 209 (1893); Luzula spadicea var. 

divaricata (S.Watson) Buchenau, in H.G.A.Engler, Pflanzenr. (iv.36) 25: 63 (1906); Luzula parviflora subsp. 

divaricata (S.Watson) Hultén, Acta Univ. Lund. (n.s.) sect. 2, 39(2): 441 (1943); Luzula parviflora var. 

divaricata (S.Watson) Boivin, Naturaliste Canad. 94: 526 (1967). T: California, Sierras, 1863, W.H.Brewer 

1794; syn: UC, fide L.Hämet-Ahti, Ann Bot. Fenn. 8: 369 (1971); W.H.Brewer 2069, 2334; syn: n.v.; Sierra 

Co., Summit Station, E.L.Greene; syn: n.v.; Sierra Co., near Castle Peak, J.G.Lemmon; syn: GH, n.v. 

Illustrations: W.L.Jepson, Fl. Calif. 1: 259, fig. 45c, d, e (1921); C.L.Hitchcock et al., Vasc. Pl. Pacif. 

Northw. 1: 216 (1969). 

Maps: L.Hämet-Ahti, Ann. Bot. Fenn. 8: 369, fig. 1 (1971); J.Coffey Swab, Fl. North Amer. 22: 258 (2000). 

Perennials (10–) 15–25 (30) cm tall, densely caespitose, pale green, subglabrous; rhizome short, 

±ascending, subnodulose; stolons absent. Basal leaves to 20 cm long, 4–6 mm wide; cauline 

leaves 2–3, 3–5 mm wide. All leaves with papillose-serrulate margins; tip acute. Lower bract 

c. 1.0–2.5 cm long, purplish-stramineous to subherbaceous, much shorter than inflorescence. 

Inflorescence many-flowered, lax, c. 7–10 cm in diam., with rigidly divaricate branches 

(often at ±right angles); flowers borne singly; distal peduncles usually more than 1.0–1.5 cm 

long. Bracteoles ovate-lanceolate, membranous; base often pale brownish, c. 1.1–1.4 mm 

long, subglabrous. Tepals equal, glabrous, stramineous to pale brown, often with reddish 

tinge, ±lanceolate, long-acuminate, (1.8–) 2.0–2.2 (–2.4) mm long; apex often ±reflexed. 

Stamens 6; anthers 0.4–0.6 mm long; filaments c. 0.4–0.5 mm long; style 0.2–0.4 mm long; 

stigmas 0.7–1.5 mm long. Capsule ±oblong, usually pale brown, acute, exceeding perianth; 

capsule segments 1.9–2.2 × 1.0–1.2 mm long. Seeds brown, 1.0–1.2 mm long, c. 0.6 mm 

wide; appendage indistinct, to 0.1 mm, fibrillate. 

An endemic of the SW United States, only found in the Sierra Nevada in California and 

westernmost Nevada. 76: CAL, NEV. Reported to occur in Oregon but material supporting 

the literature records not seen. Subalpine forest to alpine granitic slopes; 2100–3700 m. 

Map 37. 

76. USA, CALIFORNIA: Sierra Nevada, Mineral King, F.V.Coville & F.Funston 1542 (K); Butte Co., Lassen 

National Forest, Jonesville, Butte Ck, c. 1700 m, 19 Aug 1932, H.F.Copeland (H, PR); Fresno Co., Cirque Lake, 

C.H.Quibell & E.M.Quibell 7714 (W). NEVADA: Washoe Co., Mt Rose, 3100 m, A.A.Heller 10213 (K). 

18. Luzula piperi (Coville) M.E.Jones, Bull. Montana State Univ., Biol. Ser. 15: 22 

(1910) 

Juncoides piperi Coville, in C.V.Piper, Contr. U. S. Natl. Herb. 11: 185 (1906); Luzula wahlenbergii subsp. 

piperi (Coville) Hultén, Ark. Bot. 7(1): 33 (1968). T: USA, Washington, Okanogan Co., the Cascade Mtns, 

1897, A.D.E.Elmer 678; holo: US. 

45

46 


Juncoides major (Hook.) Piper, in C.V.Piper & R.K.Beattie, Fl. N.W. Coast 92 (1915); Luzula arcuata β 

[unranked] major Hook., Fl. Bor.-Amer. 2: 189 (1838). T: USA, Washington, Pierce Co., Mt Rainier, 

F.W.Tolmie; holo: K. 

Luzula wahlenbergii f. americana Kurz, Bot. Jahrb. Syst. 19: 411 (1894). T: USA, Alaska, Lynn Canal, 

‘Dejäh’, 1882, A.Krause & A.Krause; lecto: S, fide L.Hämet-Ahti, Ann. Bot. Fenn. 8: 377 (1971); isolecto: UPS. 

Luzula parviflora var. yezoensis Satake, in T.Nakai & M.Honda, Nov. Fl. Jap. 1: 27 (1938); Luzula parviflora 

subsp. yezoensis (Satake) Hultén, Fl. Alaska 297 (1968), nom. inval. T: Japan, Hokkaido, Prov. Ishikari, Mt 

Daisetsu, Jul 1931, Y.Satake; holo: TI; iso (fragment): KYO. 

Illustrations: L.Hämet-Ahti, Aquilo, Ser. Bot., 3: 18, fig. 2, 3, 4 (1965). 

Maps: L.Hämet-Ahti, Aquilo, Ser. Bot., 3: 18, fig. 5 (1965); L.Hämet-Ahti, Ann. Bot. Fenn. 8: 378, fig. 6 

(1971); J.Coffey Swab, Fl. North Amer. 22: 259 (2000). 

Perennials 20–30 (–35) cm tall, caespitose, bluish green, subglabrous; rhizome short or 

ascending; stolons absent. Basal leaves 5–10 cm long, 3–4 mm wide; cauline leaves 3, linear, 

to 3–7 cm long, 3–5 mm wide. All leaves with papillose-serrulate margins; tip acute. Lower 

bract brownish, ciliate, to c. 1 cm long. Inflorescence conspicuously nodding, usually 

c. 5 × 3 cm, usually 30–60-flowered; flowers borne singly. Bracteoles ±ovate, membranous, 

often ±pale brownish, to c. 1.5 mm long, ciliate. Tepals equal, glabrous, ±lanceolate, acute, 

(1.8–) 2.0–2.3 (–2.5) mm long, dark brown. Stamens 6; anthers (0.3–) 0.4–0.6 (–0.8) mm 

long; filaments c. 0.5 mm long; style 0.2–0.4 mm long; stigmas c. 1 mm long. Capsule 

oblong-ellipsoid, subacuminate to acute, ±castaneous-brown, exceeding perianth; capsule 

segments 1.8–2.2 (–2.4) × 1.0–1.2 mm. Seeds pale (yellowish) brown, narrowly ellipsoid, 

1.0–1.2 mm long, 0.5–0.6 mm wide; appendage indistinct; base sparsely fibrillate. 2n=24, 

fide L.Hämet-Ahti & V.Virrankoski, Ann. Bot. Fenn. 8: 158 (1971). Fig. 11. 

Western North America from the northernmost California, Idaho and Wyoming to Alaska; 

Kuril Islands, Russian Far East, and Hokkaido, Japan. 31: KAM, KUR, PRM. 38: JAP. 

70: ALU, ALA, YUK. 71: ABT, BRC. 73: IDA, MNT, ORE, WAS, WYO. 76: CAL. 

Snowbeds and mesic heaths in the mountains and in the north. Map 38. 

31. KURIL IS.: Paramushir, Kassivobara, 29 Sep 1946, D.Vorobev (VLA); Shumshu, J.Ohwi & H.Yoshii 5644 

(KYO). 38. JAPAN: Ishikari, Mt Daisetsu, Takaminega-hara, K.Inagaki 163351 (TNS). 70. ALEUTIAN 

IS.: Atka, W.J.Eyerdam 1038 (S). ALASKA: Skagway, J.P.Anderson 1646a (S); Chugach Mtns, Thompson 

Pass, L.Hämet-Ahti 860 (H). 71. ALBERTA: Banff National Park, Lake Louise, C.J.F.Skottsberg s.n. 

(UPS). BRITISH COLUMBIA: Tenquille Lake area, Crown Mtn, 1800 m, Beamish & K.Vrugtman 60980 

(OULU). 73. OREGON: Mt Hood, 29 Jul 1886, T.Howell (P). WASHINGTON: Cascade Mtns, Stevens 

Pass Region, Aug 1929, J.M.Grant (PR, S). 

19. Luzula hitchcockii Hämet-Ahti, Ann. Bot. Fenn. 8: 370 (1971) 

Luzula glabrata var. hitchcockii (Hämet-Ahti) Dorn, Vasc. Pl. Wyoming 297 (1988). T: USA, Idaho, Valley 

Co., Sawtooth Mts, Gold Fork Lookout, 8000 ft, J.W.Thompson 13769; holo WTU; iso: MICH, PR, UC, US. 

Illustrations: C.L.Hitchcock et al., Vasc. Pl. Pacif. Northw. 1: 218 (1969) as L. glabrata; L.Hämet-Ahti, Ann. 

Bot. Fenn. 8: 371, fig. 2 (1971). 

Maps: L.Hämet-Ahti, Ann. Bot. Fenn. 8: 369, fig. 1 (1971); J.Coffey Swab, Fl. North Amer. 22: 259 (2000). 

Perennials (20–) 25–30 cm tall, loosely caespitose, bright green; rhizome branched, 

creeping-ascending; stolons usually ascending. Basal leaves usually to 10 cm long, 5–6 mm 

wide; cauline leaves 3–4, 2–6 cm long, 6–8 (–10) mm wide. All leaves with papilloseserrulate 

margins; tip acute. Lower bract herbaceous, to 2.5 cm long, conspicuously shorter 

than inflorescence; middle bracts brownish or purplish membranous, ciliate. Inflorescence 

loose, ±nodding to suberect, many-flowered, c. 8 × 6 cm; flowers borne singly or less often 

in 2-flowered clusters. Bracteoles membranous, often suffused castaneous, to 1.5 mm long. 

Tepals equal, glabrous, lanceolate, (2.5–) 2.7–3.0 (–3.5) mm long, acute, dark brown to 

castaneous brown. Stamens 6; anthers (1.0–) 1.2–1.5 (–1.6) mm long; filaments 0.4–0.6 mm 

long; style 0.5–1.0 mm long; stigmas 1.5–2.5 mm long. Capsule ovoid, acute-trigonous, dark 

castaneous-brown, exceeding perianth; capsule segments (2.8–) 3.0–3.5 (–3.8) × 1.8–2.0 mm, 

with a beak to 0.9 mm long. Seeds dark brown, ±ellipsoid, c. 1.5 mm long, c. 0.7 mm wide;


Figure 11. Luzula piperi. A, habit; B, flower with bracteoles; C, seed. Luzula hitchcockii. 

D, habit; E, flower with bracteoles; F, seed. Luzula parviflora subsp. parviflora. G, habit; 

H, flower with bracteoles; I, seed. Drawn by J.R.Janish. Reproduced with permission, from 

C.L.Hitchcock, A.Cronquist, M.Ownbey & J.W.Thompson, Vasc. Pl. Pacific Northw. 1: 216 

(1969). © University of Washington Press. 

47

48 


appendage indistinct; base fibrillate. n=12 (2n=24); fide J.Pojar, Canad. J. Bot. 51: 625 

(1973). Fig. 11. 

Endemic to SW Canada (Alberta and British Columbia) and Pacific NW of USA, from 

Washington and Montana to Oregon. 71: ABT, BRC. 73: IDA, MNT, ORE, WAS. Montane 

coniferous woodlands near timberline to subalpine and alpine slopes, ridges, meadows. 

Map 39. 

71. BRITISH COLUMBIA: Manning Park, J.A.Calder & D.B.O.Savile 11642 (DAO, WTU). 73. MONTANA: 

Glacier National Park, B.Maguire & Piranian 15397 (WTU, UC). OREGON: Crater Lake National Park, 

Baker 6124 (UC). WASHINGTON: Mt Paddo, 5 Sep 1904, W.Suksdorf 4309 (L). Not located: 'from Fort 

Colville to Rocky Mts., 49° N', 1861, D.Lyall (P). 

20. Luzula glabrata (Hoppe) Desv., J. Bot. (Desvaux) 1: 143 (1808) 

Juncus glabratus Hoppe, Herb. Viv. Pl. Rar. Praes. Alp., cent. 3, sine no. (1800); Juncus intermedius Host, 

Icon. Descr. Gram. Austriac. 3: 65 (1805), nom. illeg.; Luzula intermedia [Host] Baumg., Enum. Stirp. 

Transsilv. 3: 330 (1816), nom. illeg., non Hornem. (1815); Luzula spadicea var. glabrata (Hoppe) E.Mey., 

Syn. Luzul. 8 (1823); Luzula parviflora var. intermedia [Host] Wahlenb., Fl. Suec. 217 (1824), nom. illeg.; 

Luzula glabrata var. vera Buchenau, Krit. Verz. Juncac. 83 (1879), nom. inval.; Juncoides glabrata (Hoppe) 

E.Sheld., Bull. Geol. Nat. Hist. Surv. 9: 63 (1894). T: Austria, Salzburg, Untersberg, D.H.Hoppe; lecto: 

MW, fide L.Hämet-Ahti, Ann. Bot. Fenn. 8: 371 (1971); isolecto: PR. 

Juncus pilosus [Abart] C. glabrescens Schrank, Baier. Fl. 1: 621 (1789); Luzula glabrescens (Schrank) Dalla 

Torre & Sarnth., Farn- u. Blütenpfl. Tirol. 1: 434 (1906), nom. superfl. T: [Austria] 'aus dem 

Salzburgischen [Alpen]', K.M.Moll; syn: n.v. 

Illustrations: N.T.Host, Icon. Descr. Gram. Austriac. 3: plate 99 (1805); S.Pignatti, Fl. Ital. 3: 445 (1982). 

Perennials, usually 15–30 cm tall, loosely caespitose, subglabrous to ±glabrous; rhizome 

creeping to ascending; stolons usually ascending. Leaves of sterile shoots usually 10–15 cm 

long, (4–) 5–7 (–10) mm wide; leaves at stem base usually short, 2–5 cm long; cauline leaves 

3–4, usually 3.5–6.0 cm long, 4.5–8.0 mm wide. All leaves with papillose-serrulate margins; 

apex acute. Lower bract ±herbaceous, to 2–3 cm long, much shorter than inflorescence. 

Inflorescence ±loose, ±nodding to suberect, ±divaricate, usually 5–9 × 3–6 cm, usually more 

than 35–50-flowered; flowers usually borne singly. Bracteoles ±ovate, membranous, often 

suffused brownish, c. 1.5–2.0 mm long, ciliate. Tepals equal, glabrous, ±lanceolate, 

acuminate, 2.4–3.0 mm long, deep to dark castaneous-brown, with margins indistinct or 

narrow, paler brownish. Stamens 6; anthers (1.1–) 1.4–1.6 (–2.1) mm long, up to 10 times as 

long as filaments; filaments 0.2–0.3 mm long; style 0.5–0.9 mm long; stigmas 1.0–1.5 mm 

long. Capsule ±ovoid, acute, shortly rostrate, dark brown, usually slightly exceeding 

perianth; capsule segments 2.5–2.9 × c. 1.4–1.6 mm. Seeds ±oblong-ovoid, c. 1.2–1.4 mm 

long, c. 0.7 mm wide; appendage indistinct; base fibrillate. 2n=12 [AL], fide 


Confined to the E Alps, from Berchtesgadener Alpen, Germany, to N Italy. 11: AUT, GER. 

13: ITA. Alpine grasslands, stony slopes, usually above (1700–) 2000 m. Map 40. 

11. AUSTRIA: Niederösterreich, Raxalpe, 26 Jul 1921, G.E. Du Rietz (UPS); Niederösterreich, Schneeberg, 

1800–2050 m, G.Beck [Fl. Exs. Austro-Hung.] 1871/II (H, PR, SAM, W, WU); N Salzburg, Lofer, A.Spitzel 

[H.G.Reichenbach, Fl. Germ. Exs.] 1126 (H, K, PR). GERMANY: Bavaria, Berchtesgaden, Hohe Göll, Aug 

1895, T.Wulff (LD); Berchtesgadener Alpen, Aug 1853, A.Progel (PR). 

21. Luzula desvauxii Kunth, Enum. Pl. 3: 304 (1841) 

Luzula spadicea var. desvauxii (Kunth) E.Mey., Linnaea 22: 400 (1849); Luzula glabrata var. desvauxii 

(Kunth) Buchenau, Krit. Verz. Juncac. 83 (1879); Luzula glabrata subsp. desvauxii (Kunth) K.Richt., 

Pl. Eur. 1: 185 (1890); Luzula glabrata proles desvauxii (Kunth) P.Fourn., Quatre Fl. France 153 (1935). 

T: France, Auvergne, Mont Dore, Maire; holo: B, destroyed. 

Perennials usually (25–) 35–40 (–65) cm tall, loosely caespitose, subglabrous; rhizome 

creeping, branched; stolons short, ascending. Leaves of sterile shoots usually 15–20 cm long, 

3–6 mm wide; basal leaves few, intermediate between cataphylls and cauline ones; cauline 

leaves 3–5, usually 10–20 cm long, 3.0–5.5 (–7.5) mm wide, conspicuously overlapping, 

with upper ones sometimes exceeding inflorescence. All leaves almost glabrous, with


papillose-serrulate margins; apex acute. Lower bract herbaceous, usually 2.5–5 cm long, 

shorter than or almost equalling inflorescence. Inflorescence many-flowered, ±nodding, later 

suberect, usually 5–7 × 4–5 cm; flowers borne singly or crowded in 2 (3–4)-flowered groups. 

Bracteoles scarious, ovate, ciliate, c. 1.5 mm long. Tepals equal, glabrous, lanceolate, 

acuminate, usually 2.4–2.8 mm long, deep to dark castaneous-brown; margins inconspicuous, 

paler (straw-) brown. Stamens 6; anthers (1.0–) 1.1–1.4 (–1.7) mm long; filaments 0.3–0.5 mm 

long; style c. 0.8–1.0 mm long; stigmas 1.0–1.5 mm long. Capsule oblong-ovoid, ±acute, 

shortly mucronate, dark brown, shining, ±equalling, or slightly longer than perianth; capsule 

segments (2.4–) 2.5–2.8 (–3.0) × c. 1.3–1.5 mm. Seeds ±ovoid, 1.1–1.3 mm long, c. 0.8 mm 

wide, brown; appendage to 0.1 mm long, fibrillate. 2n=12 [AL], fide A.-M.Lambert & 

J.Giesi, Inform. Annuelles Caryosyst. Cytogén. 1: 28 (1967). 

SW Europe, disjunctly in the E Pyrenees, Massif Central, Vosges and S Schwarzwald. 

Reported from Montes Cantábricos, N Spain, but material not seen [see P.Montserrat, Anales 

Inst. Bot. Cavanilles 21(2): 459-461 (1964)]. 11: GER. 12: FRA, SPA. Subalpine stony 

meadows, along streams, usually on non-calcareous substrates above 1200 m. Map 41. 

11. GERMANY: Schwarzwald, Belchen, Jul 1858, H.A. de Bary (W). 12. FRANCE: Vosges, Hohneck, 

I.S.Perrin [F.W.Schultz, Herb. Norm.] 1154 (PR); Puy-de-Dôme, Mont-Dore, Grande Cascade, 1200–1300 m, 

17 & 22 Jun 1971, Rastetter (H, LD); Auvergne, Puy-Mary, 1200 m, 28 Jul 1909, J.J.Héribaud (PR, W); 

Auvergne, Puy-Ferrand, 2 Sep 1852, E.Desvaux (H); Hautes Cévennes, S of Mt Mézenc, 1400 m, 

N.Y.Sandwith 3897 (K); E Pyrenees, Vallée d'Eyne, c. 2100 m, E.Wängsjö & G.Wängsjö 3366 (LD); E.M.Sennen 

[Pl. Esp.] 4055 (LD); E Pyrenees, L'Hospitalet, Ellman & N.Y.Sandwith 247 (K). SPAIN: E Pyrenees, 

Nuria, 1850 m, E.M.Sennen [Pl. Esp.] 1809 (BC, W). 

22. Luzula parviflora (Ehrh.) Desv., J. Bot. (Desvaux) 1: 144 (1808) 

Juncus parviflorus Ehrh., Beitr. Naturk. 6: 139 (1791); Luzula spadicea var. parviflora (Ehrh.) E.Mey., 

Linnaea 22: 402 (1849); Juncoides spadicea var. parviflora (Ehrh.) Kuntze, Revis. Gen. Pl. 2: 724 (1891); 

Juncoides parviflora (Ehrh.) Coville, Contr. U.S. Natl. Herb. 4: 209 (1893). T: [Sweden, Lappland], 

C.Linnaeus; lecto: Flora Lapponica 124, Institut de France, Paris, designated here; photo BM, PRA. 

Luzula parviflora var. densiflora Lange, Consp. Fl. Groenland. 125 (1880). T: [not indicated; numerous 

syntypes from E and W Greenland]. 

Luzula parviflora var. sparsiflora Lange, Consp. Fl. Groenland. 125 (1880). T: [not indicated; numerous 

syntypes from Greenland]. 

Map: E.Hultén & M.Fries, Atlas N. Eur. Vasc. Pl. 1: map 166 (1986). 

Perennials 30–90 cm tall, loosely caespitose; rhizome ascending; stolons short, ascending, 

less often absent. Basal leaves 10–20 cm long, usually 5–10 mm wide; cauline leaves (3–) 4– 

5, 3–10 mm wide. All leaves with papillose-serrulate margins; apex acute. Lower bract 

herbaceous or brownish membranous, usually 1.5–4.0 cm long, shorter than inflorescence. 

Inflorescence usually lax, sometimes nodding, with some main branches to 13 cm long, often 

arcuate; flowers borne singly. Bracteoles subglabrous to sparsely ciliate, ovate, membranous. 

Tepals equal, glabrous, ±lanceolate, 1.8–2.3 (–2.5) mm long. Stamens 6; anthers 0.3–0.6 mm 

long, as long as or shorter than filaments. Capsule smooth above. Seeds ±ellipsoid, brown to 

dark brown, 1.0–1.4 mm long; base fibrillate. 

A widely distributed northern circumpolar species, most common in subarctic areas and in the 

mountains of N Europe, N Asia and temperate North America. Three subspecies are recognized. 

1 Inflorescence many-flowered with long main branches and short most 

distal secondary branches (peduncles); perianth segments medium brown 22a. subsp. parviflora 

1: Inflorescence few-flowered, with long main branches and long distal 

secondary branches (almost divaricate); perianth segments usually pale 

brown to stramineous 

2 Stem conspicuously reddish at base; cauline leaves mid-green, shining, 

tepals with a distinct midrib 22b. subsp. fastigiata 

2: Stem without reddish coloration or with indistinct reddish tinge at 

base; cauline leaves bluish or greyish green, dull, tepals with an 

indistinct midrib 22c. subsp. melanocarpa 

49

50 


22a. Luzula parviflora (Ehrh.) Desv. subsp. parviflora 

Luzula neglecta Kunth, Enum. Pl. 3: 301 (1841); Luzula spadicea f. parviflora (Ehrh.) I.Grint., Fl. Rep. Soc. 

Roman. 11: 594 (1966). T: Norway, Dovrefjeld, Vollasoe, 1828, [anonymous]; syn: herb. Buek, n.v.; syn: 

‘in herb. Lucaeno’, n.v.; iso: PR. 

Illustration: J. Lid, Norsk & Svensk Fl. 202, fig. 99c (1963). 

Map: N.K.Kovtonyuk, in L.I.Malyshev & G.A.Peshkova, Fl. Sibiri 4 (Araceae – Orchidaceae): map 38 (1987) 

[Siberia]. 

Plants usually (30–) 40–60 cm tall, slightly reddish at base. Basal leaves 12–17 cm long, 

5–8 mm wide; cauline leaves (3) 4–5, lanceolate, acuminate, up to 7–9 cm long, 3–8 mm 

wide, often shorter than internodes. Inflorescence many-flowered; main branches to 9 (–10) cm 

long, nodding to suberect; distal secondary branches short (partial inflorescences not 

distinctly loose). Bracteoles entire and glabrous to sparsely ciliate. Tepals acute, not rigid; 

midrib not distinct, usually 1.9–2.3 (–2.5) mm long. Anthers 0.4–0.5 mm long; filaments 

c. 0.5 mm long; style 0.2–0.3 mm long; stigmas c. 0.5–0.7 mm long. Capsule oblong-ellipsoid, 

acute to acuminate-trigonous; mucro c. 0.2 mm long, dark brown; capsule segments 1.9–2.3 × 

1.0–1.3 mm. Seeds mid-brown, 1.1–1.3 mm long, 0.6–0.7 mm wide; appendage indistinct, to 

0.1 mm; base strongly fibrillate. 2n=24 [BL], fide H.Nordenskiöld, Hereditas 37: 330–331 

(1951); fide T.Engelskjøn, Opera Bot. 52: 14 (1979); 2n=22 [2AL+20BL], fide L.Hämet-Ahti 

& Virrankoski, Ann. Bot. Fenn. 8: 158 (1971). Fig. 11. 

N Europe, N Asia and mountains of C Asia, and northern part of North America. 10: FIN, 

NOR, SWE. 14: RUN. 30: ALT, BRY, CTA, IRK, KRA, TVA, WSB, YAK. 31: AMU, 

KAM, KUR, MAG, PRM. 32: KAZ. 36: CHN, CHX. 37: MON. 70: ASK, GNL, NWT, YUK. 

71: ABT, BRC. 72: NFL, QUE. 73: COL, IDA, ORE, WYO. 76: ARI, CAL, NEV, UTA. 

Found in a variety of habitats, from humid woodlands, meadows and stream banks, to subalpine 

grasslands and tundra. Map 42. 

10. SWEDEN: Abisko, Torneträsk Lake, 18 Jun 1948, A.Pilát & J.A.Nannfeldt (PR). 14. RUSSIA NORTH: 

Vorkuta, Aug 1983, V.V.Morozov (MW). 30. KRASNOYARSK: Yenisei R., Myelnitsa, 12 Jul 1876, 

M.Brenner (K). 31. MAGADAN: W Chukotka, Bilibino, V.Petrovskiy & T.Koroleva (VLA). 

32. KAZAKHSTAN: Kungei-Alatau, Cilik R., 23 Aug 1934, M.Popov (AA). 36. CHINA NORTH- 

CENTRAL: Shanxi, Wu-tai-shan, A.Serre A524 (UPS). 37. MONGOLIA: Bulganskiy Aimag, I.Gubanov 

9783 (MW). 70. ALASKA: Kaolak R., H.Sherman 5859 (H). GREENLAND: Nigerdleq, P.M.Jørgensen 

& Larsson [Pl. Vasc. Groenl. Exs.] 402 (C, PR). 71. BRITISH COLUMBIA: Smithers, 3–8 Aug 1931, 

V.Kujala & A.Cajander (H). 

Occurrence in Inner Mongolia, N China, is probable but material has not been seen. A very 

variable subspecies; many aberrant populations are found in the S part of the North 

American range. Intermediates between the type subspecies and subsp. fastigiata have been 

reported from Alaska, British Columbia, Wyoming, California, Utah and Colorado. 

22b. Luzula parviflora subsp. fastigiata (E.Mey.) Hämet-Ahti, Ann. Bot. Fenn. 8: 374 

(1971) 

Luzula fastigiata E.Mey., Syn. Luzul. 9 (1823); Luzula melanocarpa var. fastigiata (E.Mey.) E.Mey., Linnaea 

3: 374 (1828); Luzula parviflora var. fastigiata (E.Mey.) Buchenau, Krit. Verz. Juncac. 83 (1879). 

T: Canada, British Columbia, Nootka, T.Haenke; holo: PR, cf. L.Hämet-Ahti, Ann. Bot. Fenn. 8: 374 (1971). 

Illustration: L.Hämet-Ahti, Ann. Bot. Fenn. 8: 374, fig.4 (1971). 

Maps: L.Hämet-Ahti, Ann. Bot. Fenn. 8: 375, fig. 5 (1971); L.Hämet-Ahti, Ann. Bot. Fenn. 12: 28 (1975). 

Plants usually 30–50 (–70) cm tall, distinctly reddish at base. Basal leaves to 10 cm long, to 

8 mm wide; cauline leaves 4–5, lanceolate, acuminate, usually to 10 cm long, to 6–7 mm 

wide. Inflorescence lax, up to 13 × 12 cm, usually few-flowered, sometimes ±divaricate; 

distal inflorescence branches usually long; flowers borne singly. Bracteoles entire, ciliate. 

Tepals acute, pale brown to straw-brown, ±rigid; midrib distinct, 1.8–2.2 mm long. Anthers 

0.3–0.4 (–0.5) mm long; filaments 0.5–0.6 mm long; style c. 0.2 mm long; stigmas 0.8–1.0 mm 

long. Capsule oblong-ellipsoid, acute to acuminate, with mucro 0.1–0.2 mm long, pale brown 

to straw-brown, longer than perianth; capsule segments 2.1–2.4 × 1.1–1.3 mm. Seeds dark 

brown, 1.2–1.3 mm long, 0.7–0.8 mm wide; appendage 0.1–0.2 mm, fibrillate. 2n=24 [BL], 

fide L.Hämet-Ahti, Ann. Bot. Fenn. 8: 375 (1971).


Russian Far East and Kuril Is., Aleutian Is. and from mainland Alaska to California, Utah 

and Colorado. 31: KAM, KUR, MAG, PRM. 70: ALU, ASK. 71: ABT, BRC. 73: COL, IDA, 

ORE, WAS, WYO. 76: ARI, CAL, NEV, UTA. 77: NWM. Found in humid to mesic 

woodlands. Map 43. 

31. KAMCHATKA: Karaginskiy Distr., Ossora, 4 Sep 1976, S.Kharkevicz & T.Bucz (VLA); Middle Opala R., 

E.Hultén 2452 (S). KURIL IS., Paramushir, 13 Aug 1971, T.Neczaeva (VLA); Urup, M.Tatewaki 9857 

(SAP, TI). 70. ALEUTIAN IS.: Amchitka Is., Reich & Y.M.C.McCann 390, 621 (OULU. 71. BRITISH 

COLUMBIA: Vancouver Is., Goldstream, 27 Jun 1887, J.Macoun (BM). 73. IDAHO: Latah Co., the 

Thatuna Hills, Crumarine Ck, L.A.Dillon 564 (PR, S). OREGON: 'coniferous woods', May 1882, T.Howell 

s.n. (PR). WASHINGTON: Mt Rainier, c. 1600 m, Godman 475 (BM). WYOMING: Lincoln Co., 

Jackson's Hole, E.B.Payson & L.B.Payson 2256 (H). 

There are intermediates of presumed hybrid origin between subsp. fastigiata and the type 

subspecies (see above). 

22c. Luzula parviflora subsp. melanocarpa (Michx.) Hämet-Ahti, Ann. Bot. Fenn. 8: 

377 (1971) 

Juncus melanocarpus Michx., Fl. Bor.-Amer. 1: 190 (1803); Luzula spadicea var. melanocarpa (Michx.) 

E.Mey., Linnaea 22: 403 (1849); Luzula spadicea var. aucta Schur, Enum. Pl. Transsilv. 681 (1866), nom. 

illeg.; Luzula parviflora var. melanocarpa (Michx.) A.Gray, Manual, 5th edn, 536 (1867); Juncoides 

parviflora var. melanocarpa (Michx.) E.Sheld., Minnesota Bot. Stud. 1: 63 (1894); Luzula parviflora f. 

melanocarpa (Michx.) Neuman, Sver. Fl. 663 (1901); Juncoides parviflora subsp. melanocarpa (Michx.) 

Cockerell, in F.P.Daniels, Fl. Boulder 90 (1911); Luzula spadicea f. melanocarpa (Michx.) I.Grint., Fl. Rep. 

Soc. Roman. 11: 595 (1966); Luzula parviflora subsp. melanocarpa (Michx.) Tolm., Arktic Fl. SSSR 4: 34 

(1963), nom. inval. T: ‘Hab. in montibus sinum Hudsonis inter & Canadam, juxta lacum Cygnorum’; syn: P. 

Luzula arida Steud., Syn. Pl. Glumac. 2: 293 (1855). T: Canada, Labrador, [Hopedale, 1846–1848], 

C.G.Albrecht; holo: P, fide J.Cayouette & S.J.Darbyshire, Taxon 43: 178 (1994). 

Luzula labradorica Steud., Syn. Pl. Glumac. 2: 291 (1855), nom. illeg., non Raf. 1840. T: Canada, 

Labrador, [Hopedale, 1846–1848], C.G.Albrecht; holo: P, fide J.Cayouette & S.J.Darbyshire, Taxon 43: 178 

(1994). 

Illustration: S.E.Clemants, Juncac. New York State 50 (1990). 

Map: L.Hämet-Ahti, Ann. Bot. Fenn. 12: 28 (1975). 

Plants usually 60–90 cm tall, not or slightly reddish at base. Basal leaves dull, bluish green, 

to 20 cm long, to 8–10 mm wide; cauline leaves 4–6, 8–12 cm long, (6–) 8–10 mm wide. 

Inflorescence few-flowered but large, c. 12 (–20) × 8 cm; primary branches long; secondary 

distal branches long, spreading to reflexed; flowers borne singly. Bracteoles ±glabrous to very 

sparsely ciliate. Tepals acute, rigid, brown to pale brown, often ±translucent, c. 1.8–2.1 mm 

long; midrib not distinct. Anthers 0.4–0.6 mm long; filaments c. 0.5 mm long; style c. 0.5 mm 

long; stigmas 0.8–1.3 mm long. Capsule ±ellipsoid, acute, with mucro 0.2–0.5 mm long, 

usually dark brown, longer than perianth; capsule segments 1.9–2.2 × 1.0–1.2 mm. Seeds 

dark brown, c. 1.2 mm long; appendage not distinct; base fibrillate. 2n=24 [BL], fide Á.Löve 

& D.Löve, Univ. Colorado Stud. Ser. Biol. 17: 26 (1965). 

Eastern North America, from Labrador to Alberta, Saskatchewan and Minnesota in the west, 

and to New York State in the south. 71: ABT, MAN, SAS. 72: LAB, NBR, NFL, NSC, ONT, 

PEI, QUE. 74: MIN, WIS. 75: CON, MAI, MAS, MIC, NWH, NWY, RHO, VER. 

In mountain meadows and along streams; less often in humid montane woodlands. Map 44. 

72. LABRADOR: Twin Falls, 23 Jul 1937, I.Hustieh s.n. (H). NEWFOUNDLAND: Cormack, T.Ahti 2229 (H). 

ONTARIO: Algome Distr., High Falls, C.E.Garton et al. 14499 (H). QUÉBEC: Parc des Lamentides, 

R.Cayouette & J.Cayouette 8143 (H); Ile d'Anticosti, Crique de la Chaloupe, F.Marie-Victorin & L.Rolland- 

Germain 25799 (P). 74. MINNESOTA: Grand Marais, 26 Jun 1891, coll. unknown (H). 75. NEW 

HAMPSHIRE: Mt Washington, c. 1600 m, H.Nordenskiöld s.n. (UPS); Coos Co., Dolly Copp Road, 450 m, 

D.M.Bates & W.C.Elsik 71 (UPS). VERMONT: Mt Mansfield, 22 Jun 1877, C.G.Pringle s.n. (K). 

23. Luzula effusa Buchenau, Krit. Verz. Juncac. 88 (1879) 

Juncoides spadicea Kuntze var. effusa (Buchenau) Kuntze, Revis. Gen. Pl. 2: 724 (1891). T: Sikkim, alt. 

10000 ped. [c. 3000 m], J.D.Hooker 3; syn: K. 

51

52 


Perennials (20–) 40–80 (–100) cm tall, loosely caespitose, subglabrous; rhizome creeping or 

ascending, branched; stolons usually present, short or to c. 10 cm long. Leaves with densely 

papillose-serrulate margins; apex acute; basal leaves few; cauline leaves 3–6 (upper 2 or 3 

represent leaf-like bracts). Inflorescence usually lax, usually constituting 1/3– 1/2 of plant 

height, decompound, storied; partial inflorescences subtended by leaf-like bracts; branches 

diverging at wide angles; flowers borne singly, rarely in 2 (–3)-flowered clusters. Tepals 

equal, glabrous, ±lanceolate, usually 1.8–2.8 mm long, pale reddish brown to dark 

castaneous-brown. Stamens 6; anthers 0.5–1.1 mm long; filaments 0.6–1.0 mm long; style 

usually 0.4–0.5 mm long; stigmas 1.0–1.5 mm long. Capsule obovoid to ±ellipsoid, subacute 

to acute, with mucro 0.2–0.4 mm long, distinctly papillose above when ripe, equalling or 

±longer than perianth; capsule segments c. 2.1–2.3 × 1.1–1.4 mm. Seeds deep brown, 

1.2–1.4 mm long, (0.6–) 0.7–0.8 mm wide; apex usually distinct; basal appendage indistinct, 

to 0.1 mm, fibrillate. 

A widely distributed variable complex of populations extending from E Nepal to SW China, 

Taiwan and S Malesia. The variation is centred in Sichuan and adjacent territories where two 

varieties linked by occasional intermediates are recognized. The whole group is in need of 

revision. Most common habitats include humid mountain forests, wet scrub, shady ravines 

and cliff-faces. 

Lower cauline leaves usually 4–11 mm wide; tepals ±brown to deep 

(castaneous-)brown; stem usually less than 1.5–2.0 mm in diam. 23a. var. effusa 

Lower cauline leaves usually 10–21 mm wide; tepals paler reddish brown to 

brown; stem usually 3–6 mm in diam. 23b. var. chinensis 

23a. Luzula effusa Buchenau var. effusa 

Illustrations: Muh-Tsuen Kao & C.E.Devol, in Hui-Lin Li, Fl. Taiwan 5: 155, plate 1304 (1978); H.J.Noltie, 

Fl. Bhutan 3(1): 263, fig. 25 (1994); K.F.Wu, Fl. Reipubl. Popularis Sin. 13(3): 240, plate 58 (1997). 

Plants usually to (50–) 80 cm tall, often slender; stolons short to long. Stem usually 1–2 mm 

diam. Cauline leaves usually 4–11 mm wide. Tepals ±brown to deep (castaneous-)brown, 

usually 1.8–2.8 mm long. Anthers 0.5–1.1 mm long; filaments usually 0.6–0.8 (–1.0) mm long. 

The East Himalaya from E Nepal to Manipur, N Burma, SW and C China, Taiwan, Luzon, 

Borneo, Sulawesi and New Guinea. 36: CHC, CHN, CHT. 38: TAI. 40: EHM, NEP. 

41: BMA. 42: BOR, NWG, PHI, SUL. Humid mountain woodlands, shrubby slopes and 

subalpine ravines. Map 45. 

36. TIBET: Kongbo, Doshong La, 4000 m, F.Ludlow, G.Sherriff & G.Taylor 4676a (BM, UPS). CHINA 

SOUTH-CENTRAL: Sichuan, Mt Omei [Emei], Chiu-lo Tung, 2000 m, F.T.Wang 23312 (S). CHINA 

NORTH-CENTRAL: Gansu ['Kansu orient.'], 4 Sep 1890, Potanin (K, LE). 38. TAIWAN: Taichung, 

W slope of Mt Loyehwei-shan, Shimizu & Chuang 20183 (S). 40. NEPAL: Tamur Valley, Topke Gola, 

c. 3530 m, J.D.A.Stainton 976 (UPS). BHUTAN: Pele-La, Gould 345 (K). 42. BORNEO: Sabah, 

Kinabalu, Masilau R., M.S.Clemens 51371 (K). NEW GUINEA: Papua, Goilala, Mt Albert Edward, 3600 m, 

J.R.Croft et al. 61394 (BM, K). PHILIPPINES: Luzon, Mt Pulog, E.D.Merrill 6490 (K). SULAWESI: 

Enrekang, Rante Mario, 3300 m, P.J.Eyma 879 (K). 

A very variable taxon. Rarely, very young plants have the inflorescence only indistinctly 

storied. Plants from Taiwan and Malaya have tepals usually 2.5–2.8 mm long, longer than 

capsules. The Malayan plants have more numerous basal leaves and, on average, longer 

stolons. Intermediates between the two varieties are not rare. 

23b. Luzula effusa var. chinensis (N.E.Br.) K.F.Wu, J. E. China Norm. Univ., Nat. Sci. 

Ed., 1992(3): 92 (1992) 

Luzula chinensis N.E.Br., J. Linn. Soc., Bot. 36: 161 (1903). T: China, [Sichuan] Szechuan, A.Henry 8829; 

syn: K; S (fragment); China, Szechuan, near Tibetan frontier, Tachienlu [Kangding], A.Pratt 360; syn: BM, K. 

Plants usually 50–100 cm tall, robust; stolons short. Stem usually 2–6 mm in diam. Cauline 

leaves usually to 20 cm long and 10–21 mm wide. Tepals usually 1.9–2.3 mm long, paler 

reddish brown to brown. Anthers 0.5–0.6 mm long; filaments 0.6–0.8 mm long.


Most frequently found in Sichuan and Yunnan, less common in Tibet, Hubei, Guizhou and 

N Burma. 36: CHC, CHT. 41: BMA. Mesic to humid woodlands. Map 46. 

36. CHINA SOUTH-CENTRAL: Hubei [originally Hupeh, without detailed locality], A.Henry 6809 (K); 

Sichuan, Ta-hsiang-ling, 2700 m, H.Smith 2083 (UPS); Nanchuan-hsien, c. 200–2200 m, W.P.Fang 974 (E); 

1157 (E, K); Omei [Emei] Shan, E.H.Wilson 5303, 5304 (BM, K); Yunnan, Yangbi Xian, W side of Diancang 

Shan, 3300–3600 m, B.Bartholomew et al. 423 (E). TIBET: Pome [Bomi], Tongkah, F.Ludlow, G.Sherriff & 

R.H.J.Williams 13770 (BM). 

Extremely robust forms of var. chinensis are very different from the type variety but 

intermediates are known. 

24. Luzula alpinopilosa (Chaix) Breistr., Bull. Soc. Sci. Isére 61: 609 (1947) 

Juncus alpinopilosus Chaix, in D.Villars, Hist. Pl. Dauphiné 1: 318 (1786). T: Haller 1326 [A. von Haller, 

Hist. Stirp. Helv. 2: no. 1326 (1768)]; syn: P-Haller; France, Chaliol-le-Viel, D.Chaix; syn: G, n.v. 

Luzula variabilis Buchenau, Bot. Jahrb. Syst. 7: 171 (1885), [‘species collectiva’], nom. illeg. 

Perennials 10–65 cm tall, subglabrous, loosely caespitose; rhizome branched, ascending; 

stolons often present, short. Basal leaves few, intermediate between cataphylls and cauline 

leaves; cauline leaves 3–5, 1.5–9.0 mm wide. All leaves with papillose-serrulate margins; 

apex acute. Lower bract herbaceous to subherbaceous, shorter than or rarely ±equalling 

inflorescence. Inflorescence a loose to subdense suberect or ±nodding panicle to 7 × 6 cm; 

flowers borne singly and in 2–3 (–7)-flowered clusters. Bracteoles ovate-lanceolate, 1.5–2.0 mm 

long, brownish, ciliate. Tepals ±equal, glabrous, lanceolate, with outer ones acuminate, 

1.8–3.3 mm long, blackish brown to dark castaneous-brown; margins indistinct or very 

narrow. Stamens 6; anthers 0.6–1.2 mm long, 3–9 times longer than filaments; filaments 

0.1–0.4 mm long; style 0.6–1.2 mm long; stigmas 1–2 mm long. Capsule ovoid to oblongovoid, 

subobtuse to ±acute, shortly mucronate, usually ±exceeding perianth, dark castaneousbrown, 

smooth; capsule segments usually 2.1–2.4 × 1.2–1.5 mm. Seeds pale brown, oblongovoid, 

1.1–1.3 mm long, 0.6–0.7 (–0.9) mm wide; apex distinct, pale, to 0.1–0.2 mm long; 

basal appendage minute, as a paler patch to 0.1 mm long; base fibrillate. 

Mountains of C and S Europe. Very variable in size, leaf width, tepal length and 

anther/filament length ratio. Three subspecies are recognized but numerous intermediates 

exist. Subalpine and alpine grasslands, rocky slopes, on gravel along streams, confined to 

silicic substrates. 

1 Filaments 0.1–0.2 (–0.3) mm long; anthers 4–9 times longer than 

filaments 24b. subsp. obscura 

1: Filaments 0.3–0.4 mm long; anthers usually 2–4 times longer than 

filaments 

2 Cauline leaves 4–7 cm long, 1.5–5.5 mm wide; anthers usually 

0.9–1.2 mm long; lower bract substantially shorter than inflorescence 24a. subsp. alpinopilosa 

2: Cauline leaves 7–13 cm long, 4–9 mm wide; anthers usually 

0.7–0.8 mm long; lower bract ±equalling inflorescence or shorter 24c. subsp. deflexa 

24a. Luzula alpinopilosa (Chaix) Breistr. subsp. alpinopilosa 

Juncus spadiceus All., Fl. Pedem. 2: 216 (1785), nom. illeg., non Vill. (1779); Luzula spadicea [All.] DC., Fl. 

Franç., 3rd edn, 3: 159 (1805); Luzula spadicea var. allionii E.Mey., Linnaea 22: 401 (1849), nom. inval.; 

Luzula spadicea var. barbata Neilr., Nachtr. Fl. Wien 100 (1851); Luzula variabilis Buchenau, Bot. Jahrb. 

Syst. 7: 171 (1885), [doubtfully valid: ‘species collectiva’], nom. illeg.; Juncoides spadicea Kuntze, Revis. 

Gen. Pl. 2: 724 (1891). T: Haller 1326; syn: P-Haller; Scheuchzer It. Alp. 6: 458; syn: n.v. [authentic 

material of Scheuchzer in W] 

Luzula spadicea var. laxiflora Desv., J. Bot. (Desvaux) 1: 146 (1809); Luzula spadicea f. laxiflora (Desv.) 

I.Grint., Fl. Rep. Soc. Roman. 11: 594 (1966). T: 'in pratis humidis Alpium Europae' [otherwise not 

indicated]. 

Luzula spadicea β [unranked] elatior Gaudin, Fl. Helv. 2: 569 (1828). T: [Switzerland] in valle Champé, 

J.Gay; syn: n.v. 

53

54 


Luzula spadicea var. candollei E.Mey., Linnaea 22: 401 (1849); Luzula spadicea subsp. candollei (E.Mey.) 

K.Richt., Pl. Eur. 1: 185 (1890); Luzula alpinopilosa subsp. candollei (E.Mey.) Rothm., Feddes Repert. Spec. 

Nov. Regni Veg. 67: 4 (1963). T: France, E Pyrenees, Val d’Eynes, 13 Sep 1839, R.J.Shuttleworth; syn: W; 

Switzerland, Geneve, Castan; syn: n.v.; Pyrenees, Meles, L.Marchand; syn: n.v. 

Illustrations: H.E.Hess, E.Landolt & R.Hirzel, Fl. Schweiz 1: 519 (1967), as L. spadicea; S.Pignatti, Fl. Ital. 3: 

445 (1982). 

Plants (10–) 15–20 (–35) cm tall. Leaves of sterile shoots (when present) usually 5–10 (–20) 

cm long, 2–4 (–5) mm wide; cauline leaves usually 4–7 cm long, 1.5–3.5 (–5.5) mm wide. 

Lower bracts subherbaceous or pale brownish, 2–3 (–4) cm long, substantially shorter than 

inflorescence. Inflorescence usually 4–7 × 3–6 cm, suberect to slightly nodding, ±evenly 

branched, most flowers usually borne singly, but with 2–4-flowered clusters usually also 

present. Tepals 1.8–2.7 mm long. Anthers 0.9–1.2 mm long, usually 3–4 times longer than 

filaments; filaments 0.3–0.4 mm long; style (0.7–) 0.9–1.2 mm long; stigmas c. 1.5 mm long. 

2n=12 [AL], fide H.Nordenskiöld, Hereditas 37: 330–331 (1951); Druskovic, Int. Organ. Pl. 

Biosyst. Newslett. 24: 13 (1995). 

Mountains of Central and southern Europe: the Alps, the N Apennines, the Pyrenees, the 

Vosges. 11: AUT, GER, SWI. 12: FRA, SPA. 13: ITA, YUG-SL. Rocky and stony subalpine 

and alpine slopes. Map 47. 

11. AUSTRIA: Steiermark, Bösenstein, 1896 m, G.Strobl [Fl. Exs. Austro-Hung.] 1872 (PR, PRC, WU); 

Vorarlberg, Montafon, Hochmaderjoch, Gölles 462 (WU). SWITZERLAND: Tessin, St. Gotthard, Monte 

Prosa, 2400 m, A.Kneucker [Cyper. Junc. Exs.] 90 (PR, PRA); Valais, Zermatt, Riffelberg, F.O.Wolf 

[A.Kneucker, Cyper. Junc. Exs.] 90a (PR). 13. ITALY: N Apennines, Corno alle Scale, 21 Jul 1840, P.Savi 

(PR); Saint-Rhémy, 2400 m, A.Charpin, W.Greuter & Hainard [Exs. Genav.] 117 (W). 

Very variable but the two extremes (the broad-leaved, small-flowered form often recognized 

as subsp. candollei, and the typical narrow-leaved large flowered plants) often coexist within 

populations, and intermediates are not rare. 

24b. Luzula alpinopilosa subsp. obscura S.E.Fröhner, Preslia 40: 426 (1968) 

Luzula obscura (S.E.Fröhner) Novikov, Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 95(6): 66 (1990). 

T: Slovakia, Vysoké Tatry, Zbojnická chata, 1900 m, 14 Aug 1965, S.Fröhner; holo: PR. 

Luzula carpatica Kitt. ex Kanitz, Linnaea 32: 327 (1863), an nom. validum?; Luzula spadicea A.b. [unranked] 

carpatica (Kitt. ex Kanitz) Asch. & Graebn., Syn. Mitteleur. Fl. 2(2): 513 (1904);. Luzula spadicea f. 

carpatica (Kitt. ex Kanitz) I.Grint., Fl. Rep. Soc. Roman. 11: 594 (1966). T: [Slovakia], 'in der kleinen 

Kohlbach', J.D.T.Mauksch; holo: n.v., probably BP. 

Illustration: T.Szynal & J.Mądalski, Atlas Fl. Polsk. 2(2): 151 (1931). 

Plants 10–30 cm tall. Cauline leaves 3–8 cm long, 3.5–6.5 mm wide. Lower bract usually 

herbaceous, usually 1.5–4.5 cm long, shorter than inflorescence. Inflorescence c. 4–5 × 3–5 cm, 

±nodding to suberect; main branches long; secondary branches often abbreviated; flowers 

borne singly and usually in (2–) 3–5 (–7)-flowered loose clusters. Tepals (2.0–) 2.2–3.0 mm long. 

Anthers 0.9–1.2 mm long, usually 4–9 times longer than filaments; filaments 0.1–0.2 (–0.3) mm 

long; style 0.8–0.9 mm long; stigmas 1.2–2.0 mm long. 2n=12 [AL], fide J.Májovský & 

A.Murín, Karyotax. Prehl. Fl. Slov. 361 (1987). Fig. 12. 

C Europe, confined to the Carpathians from Slovakia and Poland to Romania. 11: CZE-SL, 

POL. 13: ROM. 14: UKR. Map 48. 

11. SLOVAKIA: Nízké Tatry, Králova Hola, 1700 m, 20 Aug 1898, K.Tocl (PR); Vysoké Tatry, Mlynická 

dolina, 1700 m, 10 Jul 1925, K.H.Rechinger (W). 13. ROMANIA: Retezat [Mtns], Gura Apei, Zanoga Lake, 

2 Jul 1970, J.Soják (PR); Muntii Fogarasului, Vistea, Sambatei Valley, 1200–2000 m, 30 Jun 1970, J.Soják 

(PR). 14. UKRAINE: the East Carpathians, Mt Goverla, 2030 m, 7 Jul 1948, E.Bradis & A.Zanjatova 

(MW); Zakarpatskaya area, Rachiv, Mt Turkul (1850 m) 16 km NE of Bogdan, 1740 m, 27 Jul 1996, Z.Kaplan 

96/506 (PRA). 

The overall variation of subsp. obscura is greater than that described in the protologue. 

In particular, narrow-leaved and small-flowered plants occur in populations of subsp. 

obscura. Some plants from Bulgaria (the Stara Planina Mtns) are intermediate between 

subsp. obscura and subsp. deflexa.


Figure 12. Luzula alpinopilosa subsp. obscura. A, habit; B, inflorescence; C, terminal group 

of flowers; D, flower; E, capsule; F, seed; G, tepal; H, bracteole; I, stem T.S.; J, leaf T.S. 

(A, C–J, Piasecki, 4 Aug 1890, KRA; B, A.Rehman s.n., KRA). Scale bars: A, B = 5 cm; 

C = 5 mm; D–H = 2 mm; I, J = 1 mm. Drawn by T.Szynal. Reproduced with permission, 

from T.Szynal & J.Mądalski, Atlas Fl. Pol. 2/2: 151 (1931). 

55

56 


24c. Luzula alpinopilosa subsp. deflexa (Kožuharov) Kirschner, Taxon 50: 1109 (2001) 

Luzula deflexa Kožuharov, Fl. Bulg. 2: 402 (1965); Luzula glabrata subsp. deflexa (Kozhukh.) Kozhukh., in 

N.Andreev et al., Opred. Vissh. Rast. B'lgar. 786 (1992). T: Bulgaria, [Mtns] Vitosha, A.Yavashev; holo: 

SOM. 

Luzula spadicea var. sarplaninae Rohlena, Věstn. Král. České Společn. Nauk, Tr. Mat-Prir 1937(1)6: 7 (1938). 

T: F.Y.R. Macedonia, Šar Planina, Crni vrh, 2400–2500 m, 14 Jul 1934, K.Hrubý, A.Jirásek & T.Martinec; 

holo: PRC. 

Luzula spadicea var. velenovskyi Kožuharov, Izv. Bot. Inst. Bulg. Akad. (Sofia) 11: 130 (1963), nom. inval.; 

Luzula alpinopilosa subsp. velenovskyi [Kožuharov ex] Chrtek & Křísa, Preslia 46: 212 (1974), nom. inval. 

Authentic material: Bulgaria, 'Rila, Rodopi', V.Stříbrný; PRC, SOM. 

Luzula spadicea var. davidovii Kožuharov, Izv. Bot. Inst. (Sofia) 11: 130 (1963), nom. inval. Authentic 

material: Bulgaria, Vitosha, A.Yavashev; SOM. 

Luzula spadicea f. pumila Kožuharov, in D.Jordanov, Fl. (N. R.) B'lgar. 2: 402 (1964), an nom. valid.? 

T: Bulgaria, Mt Rila reg. alpina [collector not given]; holo: n.v. 

Luzula spadicea f. compacta Kožuharov, in D.Jordanov, Fl. (N. R.) B'lgar. 2: 402 (1964), an nom. valid.? 

T: Bulgaria, Mt Pirin reg. alpina [collector not given]; holo: n.v. 

Plants (10–) 25–50 (–65) cm tall. Leaves of sterile shoots (when present) to 30 cm long, 

c. 5–7 mm wide; cauline leaves usually 4–5, usually (7–) 9–13 cm long, 4–8 (–9) mm wide. 

Lower bract herbaceous, usually (2–) 4–8 cm long, ±equalling or shorter than inflorescence. 

Inflorescence usually 4–7 × 3–6 cm; main branches long; secondary branches short; flowers 

usually in 2–3 (–5)-flowered clusters, some borne singly. Tepals (2.0–) 2.5–2.8 (–3.3) mm 

long. Anthers 0.7–0.8 (–1.0) mm long, 2–3 times longer than filaments; filaments 0.3–0.4 mm 

long; style 0.6–0.7 mm long; stigmas 1.0–2.0 mm long. 

S Europe, confined to Balkan Peninsula: Albania, Bulgaria, F.Y.R. Macedonia and Greece. 

14: ALB, BUL, GRC, YUC. Subalpine and alpine grasslands, wet scree, rocky slopes, rock 

ledges. Map 49. 

14. BULGARIA: the Rila [Mtns], Malyovitsa, near Malyovitsa Chalet, c. 1800 m, J.Kirschner B-664 (PRA); 

the Rila [Mtns], Mt Polič, 27 Jul 1912, B.Davidov (SOM); the Rila [Mtns], Mermera Mtn, 2630 m, 5 Aug 

1964, M.Simeonovski (SOM); the Pirin, Todorin Vrkh, 14 Jul 1926, D.Yordanov (SOM). F.Y.R. 

MACEDONIA: Shar Planina, above Shtrbtse, 1900 m, K.Vandas (PR). 

Very variable in size, tepal length and leaf width. In many characters it overlaps with those 

of subsp. obscura. Plants intermediate between subsp. obscura and subsp. deflexa occur in 

N Bulgaria; plants from Greece probably are similar. 

Presumed hybrids 

Luzula parviflora (Ehrh.) Desv. × L. piperi (Coville) M.E.Jones 

Reputed hybrid, fide L.Hämet-Ahti, Ann. Bot. Fenn. 8: 378 (1971). The only specimen 

referred to the hybrid combination, [USA, Montana, Ravalli Co., C.L.Hitchcock & 

C.V.Muhlick 15340 (UC)] is reported to be seed sterile. 

Luzula × sichuanensis K.F.Wu, J. E. China Norm. Univ., Nat. Sci. Ed., 1992(3): 92 (1992), 

pro sp. 

T: China, Sichuan, Maerkang Xian, Ma-tang, H.Smith 4385; holo: PE, photo TI, PRA; iso: UPS. 

Plants to 50–60 cm tall; rhizome short, ascending; stolons short. Basal leaves to 25 cm long, 

4–6 mm wide; cauline leaves (1–) 2–3 (–4), 8–12 cm long, 4–6.5 mm wide. All leaves 

subglabrous, with papillose-serrulate margins; apex acute. Inflorescence lax, to 12 × 12 cm, 

terminal (not storied), few-flowered; flowers borne singly; secondary branches long. 

Bracteoles ±entire, later sublacerate above. Tepals ±equal, lanceolate, (1.5–) 1.7–1.9 mm 

long, mid-brown to paler brown. Stamens 6; anthers 0.3–0.5 mm long (in the UPS isotype; 

original description gives 0.6–0.8 mm); filaments 0.4–0.5 mm long; style c. 0.4 mm long; 

stigmas 1.2–1.5 mm long. Capsule ±ovoid-ellipsoid, subacute, longer than perianth, dark


castaneous-brown, smooth above; capsule segments 1.9–2.1 × 1.3–1.4 mm. Seeds 1.2–1.3 mm 

long, c. 0.7 mm wide; appendage indistinct; base fibrillate. 

The type plant is intermediate between L. parviflora and L. effusa in many characters. The 

material is too scanty to allow a more definite conclusion; it may be a result of hybridization 

or old introgression, or alternatively, it may represent a local, isolated aberrant derived from 

L. parviflora. In addition to the above locality, similar plants but closer to L. effusa were 

identified from SW Sichuan, near Muli monastery, H.Handel-Mazzetti 7475 (WU); they are 

more robust and have slightly papillose capsules. All these intermediates seem to be fertile. 

Sect. 5. ALPINAE 

Luzula sect. Alpinae Chrtek & Křísa, Preslia 46: 211 (1974) 

Luzula subsect. Alpinae (Chrtek & Křísa) Kovtonyuk, Bot. Žurn. (Moscow & Leningrad) 72: 1399 (1987); 

Luzula sect. Spicatae K.F.Wu, J. E. China Norm. Univ., Nat. Sci. Ed., 1992(3): 101 (1992), nom. illeg. 

T: Luzula spicata (L.) DC. 

Perennials. Leaves with acute to acuminate tips (mostly all leaves, sometimes only cauline 

ones). Lower bract herbaceous to membranous. Inflorescence usually nodding, less often 

erect (if so, then usually a ±compact single head), spike-like or paniculate, congested or 

interrupted at base. Seeds subglobular to ellipsoid; appendage indistinct, to 0.2 mm long or 

±lacking; base fibrillate. Diploids or agmatoploids. 

An intricate, not yet fully monographed group of 20 species distributed in the mountains of 

the northern hemisphere, and in South America and New Zealand, again mainly in high 

mountains. Here a new revision is presented but a number of questions remain unanswered. 

1 Abaxial surface of basal leaves with thick white tomentum 25. L. ulophylla 

1: Abaxial surface of basal leaves glabrous (margins usually ciliate) 

2 Plants stoloniferous 

3 Stem 10–45 cm long; inflorescence ±nodding; stem much overtopping 

basal leaves; tepals reddish-blackish 42. L. vulcanica 

3: Stem less than 5 cm long; inflorescence ±erect; stem shorter than or 

scarcely equalling basal leaves, often not visible; tepals whitishmembranous 

with pale brown midrib 27. L. celata 

2: Stolons absent 

4 Stamens 3, rarely 4–5 

5 Inflorescence an interrupted panicle composed of remote panicles of 

spike-like flower clusters; inflorescence constitutes at least 1/3 of 

plant height 

6 Outer tepals usually 2.5–3.0 mm long; style 0.4–0.5 mm long; 

stigmas 1.0–1.2 mm long 32. L. ecuadoriensis 

6: Outer tepals usually up to 2 mm long; style 0.2–0.4 mm long; 

stigmas 0.5–0.7 mm long 33. L. excelsa 

5: Inflorescence ±spike-like (not composed of panicles of clusters), 

compact or interrupted or of usually 3–6 cylindrical ±pedunculate 

clusters; inflorescence constitutes less than 1/5 of plant height 

7 Tepals ciliate 

8 Plants to 13 cm tall; stem thin, c. 0.3–0.4 mm diam. in upper part; 

inflorescence ±erect to slightly nodding, usually 2.5–5 cm long 43. L. antarctica 

8: Plants usually 20–60 cm tall; stem rigid, about 1 mm diam. in 

upper part; inflorescence nodding, usually 1.0–1.5 cm long 41. L. peruviana 

57

58 


7: Tepals ±glabrous (not to be confused with ciliate bracteoles) 

9 Inflorescence usually distinctly nodding 

10 Style 0.1–0.2 mm long; capsule segments 1.5–1.8 mm long; leaf 

margin ±densely papillose-serrulate 40. L. racemosa 

10: Style 0.3–0.4 mm long; capsule segments 1.8–2.1 mm long; leaf 

margin ±smooth 26. L. traversii 

9: Inflorescence erect to suberect 

11 Inflorescence composed of 3–6 (10) distinct elongated 

cylindrical flower clusters; lower clusters pedunculate and 

upper subsessile 37. L. brachyphylla 

11: Inflorescence congested, usually a single head, ovoid to lobate, 

rarely interrupted at base; clusters ±sessile 

12 Basal leaves 4–9 mm wide; basal bract herbaceous; style 

c. 0.3 mm long. [Juan Fernández Is.] 35. L. masafuerana 

12: Basal leaves c. 1 mm wide; basal bract membranous; style 

c. 0.1 mm long. [NW Argentina] 44. L. ruiz-lealii 

4: Stamens 6, rarely 4 or 5 

13 Tepals densely to sparsely ciliate 36. L. alopecurus 

13: Tepals ±glabrous 

14 Seeds (excluding basal appendage) more than 1.2 mm long 

15 Seeds 2.1–2.4 mm long 30. L. pediformis 

15: Seeds less than 2.0 mm long 

16 Anthers 2.0–2.9 mm long, at least 5 times as long as filaments 31. L. caespitosa 

16: Anthers less than 1.2 (–1.5) mm, up to 3 times as long as 

filaments 

17 Style 0.6–0.9 mm. [S Europe] 28. L. pindica 

17: Style 0.3–0.6 mm. [South America] 

18 Tepals ±equal, c. 2.7–3.3 mm long; anthers usually 

0.6–0.9 mm long; inflorescence ±compact 38. L. chilensis 

18: Tepals unequal, with outer ones 3.5–4.8 mm long; anthers 

usually 0.7–1.5 mm long; inflorescence often interrupted 

at base 39. L. leiboldii 

14: Seeds (excluding basal appendage) less than 1.2 mm long 

19 Inflorescence erect to suberect 

20 Inflorescence of a single compact ±ovoid head; cauline 

leaves absent; basal leaves ±obtuse; basal bract membranous, 

to 1 cm long 44. L. ruiz-lealii 

20: Inflorescence composed of 3–6 (–10) distinct spike-like 

elongated cylindrical flower clusters; lower clusters pedunculate 

and upper ones subsessile; cauline leaves 1 or 2; basal bract 

herbaceous, 1.5–4.0 cm long 37. L. brachyphylla 

19: Inflorescence nodding 

21 Inflorescence a panicle of remotely interrupted partial 

inflorescences, each of long pedunculate secondary panicles 

composed of 3–10 spike-like clusters; lowermost secondary 

panicle often 10–15 cm below the terminal one 34. L. mendocina 

21: Inflorescence congested in a single head, sometimes lobate or 

interrupted at base; basal clusters subsessile, not remote


22 Leaf margin ±smooth; basal leaves acute with acicular tip 

22: Leaf margin ±densely papillose-serrulate; basal leaves ±obtuse 

26. L. traversii 

to acute (on a single plant) 29. L. spicata 

25. Luzula ulophylla (Buchenau) Cockayne & Laing, Trans. Proc. New Zealand Inst. 

43: 366 (1911) 

Luzula racemosa var. ulophylla Buchenau, Oesterr. Bot. Z. 48: 245 (1898). T: New Zealand, South Island, 

Albertown, Lake Wanaka, D.Petrie; syn: W [seen in 1994, in 1998 not located]; Southern Canterbury Alps, 

Castle Hill Distr., 2200 ft., Dec 1894, L.C.Cockayne; syn: B, destroyed; Castle Hill, 1893, L.C.Cockayne; neo: 

WELT 12668, fide E.Edgar, New Zealand J. Bot. 4: 165 (1966); E.Edgar, in L.B.Moore & E.Edgar, Fl. New 

Zealand 2: 72 (1970). 

Perennials (5–) 9–20 (–28) cm long, densely hairy, densely caespitose; rhizome short, 

vertical, ±many-headed; stolons absent. Basal leaves short, usually 3–6 cm long, (1–) 2–3 mm 

wide, usually inrolled, narrow, whole, or upper two thirds of abaxial surface with thick white 

tomentum; cauline leaves absent or 1 (–2), with blade reduced, filiform, 1–2 cm long; sheath 

to 2 cm long. All leaves with subulate acute tip; margins ±smooth. Lower bract ovate, strawbrown, 

membranous, c. 5 mm long, similar to middle bracts. Inflorescence erect, cylindrical 

to ±ovoid, densely congested, spike-like, usually 0.8–2.0 × 0.6–1.0 cm, of 1–5 few-flowered 

clusters. Bracteoles ovate, scarious, 1.5–2.0 mm long, ciliate near apex. Tepals ±entire, 

subequal, rarely ±denticulate near apex, (1.3–) 1.6–1.9 (–2.1) mm long, acuminate; lower 

middle part dark castaneous brown; margins broad, white-scarious. Stamens 3 (4–6); anthers 

0.4–0.5 (–0.7) mm long; filaments 0.3–0.5 mm long; style c. 0.3 mm long; stigmas c. 0.5 mm 

long. Capsule obovoid, equalling or slightly exceeding perianth, ±obtuse to subacute, dark 

brown to black; capsule segments 1.5–2.0 × c. 1.1–1.3 mm. Seeds pale castaneous, 0.8–0.9 mm 

long, c. 0.6 mm wide; appendage minute, appearing as a paler basal fibrillate swelling to 

0.1 mm long. 2n=48 [48CL], fide H.Nordenskiöld, New Zealand J. Bot. 4: 186 (1966); 

2n=44, fide J.B.Hair, New Zealand J. Bot. 5: 18 (1967). Fig. 13. 

Endemic to the South Is. of New Zealand. 51: NZS. Wind-eroded shingly open ground or 

grasslands, between 300 and 1500 m. Map 50. 

51. NEW ZEALAND SOUTH: South Is., Otago, head of Eweburn Dam, H.E.Connor & H.Nordenskiöld S151 

(UPS); C Otago, Queenstown, Ben Lomond, H.E.Connor & H.Nordenskiöld S189 (UPS); Canterbury, Acheron 

R., near diversion to Lake Coleridge, E.Edgar G5791 (CHR, PR); S Canterbury, Lake Tekapo, Boundary 

Stream, 13 Jan 1961, H.E.Connor (CHR 113675, K); Canterbury, Lake Heron, 720 m, A.D.Wilton 98048 

(CHR, PRA). 

Easily distinguished by the dense white tomentum on the abaxial surface of the leaves. 

26. Luzula traversii (Buchenau) Cheeseman, Man. New Zealand Fl., 2nd edn, 305 (1925) 

Luzula racemosa var. traversii Buchenau, Bot. Jahrb. Syst. 12: 133 (1890). T: New Zealand, H.H.Travers; 

lecto: WELT 12718, fide E.Edgar, New Zealand J. Bot. 4: 164 (1966); E.Edgar, in L.B.Moore & E.Edgar, Fl. 

New Zealand 2: 71 (1970). 

Perennials 10–40 cm tall, densely caespitose; rhizome short; stolons absent. Leaves with 

±acicular tip and sparsely ciliate, with ±smooth margins. Lower bract herbaceous, exceeding 

inflorescence, 3–5 cm long. Inflorescence ±nodding, of (1–) 3–9 clusters congested into a lobate 

or pyramidal head up to 2–3 × 1.5 cm. Bracteoles ovate, entirely membranous, often castaneous 

at base, 1.5–2.0 mm long, ±entire or sparsely ciliate near apex. Tepals subequal, ±entire, 

usually with a dark castaneous brown midrib and broad whitish-membranous margins. Style 

0.3–0.4 mm long; stigmas c. 1.0–1.2 mm long. Capsule subacute, equalling or ±exceeding 

perianth. Seeds 1.0–1.2 mm long, 0.6–0.7 mm wide; appendage indistinct, to 0.1 mm, fibrillate. 

Endemic to the South Is. of New Zealand. Similar to L. racemosa but inflorescence not 

interrupted, tepals with broad scarious-membranous margins, and longer style, capsule 

segments and seeds. L. masafuerana differs from L. traversii in having broader leaves with 

papillose-serrulate margins, broader bracts and suberect inflorescence. Two varieties are 

recognized within L. traversii; intermediates are known (Canterbury, Cave Stream, N.T.Moar 

& H.Nordenskiöld S118, UPS). E.Edgar, New Zealand J. Bot. 4: 164 (1966), mentioned 

59

60 


Figure 13. Luzula ulophylla. A, habit; B, inflorescence; C, perianth and capsule; D, tepal 

and stamen; E, leaf apex, note the abaxial surface indumentum and acute tip; F, seeds. 

(A, B, E.Edgar G5791, PR; C–F, H.E.Connor & H.Nordenskiöld S151, UPS). Scale bars: 

A = 5 cm; B = 5 mm; C = 1 mm; D = 1 mm; E = 1 cm; F = 1 mm. Drawn by © E.Smrčinová.


dwarf plants from C Otago as a part of the variation range of L. traversii. The absence of 

herbaceous bracts might indicate a distinct taxon. 

Stamens usually 3; anthers ±as long as filaments; basal leaves usually 2–4 mm wide 26a. var. traversii 

Stamens usually 6; anther/filament ratio c. 2; basal leaves usually to 2 mm wide 26b. var. tenuis 

26a. Luzula traversii (Buchenau) Cheeseman var. traversii 

Basal leaves usually up to 12–15 cm long, (1–) 2–4 (–5) mm wide; cauline leaves 2 or 3, 

usually up to 6–7 cm long. Tepals 1.8–2.2 (–2.5) mm long. Stamens 3– (4–5); anthers 

0.5–0.6 mm long; filaments 0.4–0.5 mm long. Capsule castaneous brown; capsule segments 

1.8–1.9 × c. 1.2 mm. 2n=46 [46CL], fide H.Nordenskiöld, New Zealand J. Bot. 4: 186 

(1966); 2n=32, fide J.B.Hair, New Zealand J. Bot. 5: 18 (1967). 

Endemic to the South Is. of New Zealand. 51: NZS. Rocky slopes, 600–1500 m. Map 51. 

51. NEW ZEALAND SOUTH: South Is., Canterbury, Mt. Cook, Mueller Moraines, 800–950 m, N.T.Moar S82 

(UPS); Canterbury, Mt Cook District, Hooker Valley, J.D.Lovis 1087 (BM); Mt Cook, Bull Hut, H.St.John 

24675 (K); Otago, Lindis Pass, 1450 m, N.T.Moar & H.Nordenskiöld S225 (UPS); Broken R., A.Sinclair & 

J.F.J. von Haast 236 (K); Craigieburn Range, Mt Hamilton, R.Melville 5695C (K). 

26b. Luzula traversii var. tenuis Edgar, New Zealand J. Bot. 4: 164 (1966) 

T: New Zealand, South Island, C Otago, Cromwell Gorge, 5 Feb 1958 H.Nordenskiöld S173 & H.E.Connor; 

holo: CHR113666; iso: UPS. 

Basal leaves usually up to 10–15 cm long, 1–2 (–2.5) mm wide; cauline leaves (1–) 2 (–3), 

up to 6–7 cm long. Tepals 2.1–2.8 mm long. Stamens 6; anthers 0.8–1.0 mm long; filaments 

0.4–0.5 mm long. Capsule dark castaneous brown; capsule segments 1.8–2.1 × 1.3–1.4 mm. 

2n=46 [46CL], fide H.Nordenskiöld, New Zealand J. Bot. 4: 186 (1966); 2n=42, fide 

J.B.Hair, New Zealand J. Bot. 5: 18 (1967). 

Known only from a small area of C Otago, South Is., New Zealand. 51: NZS. Rocky slopes, 

200–450 m. Map 52. 

51. NEW ZEALAND SOUTH: South Is., C Otago, Lindis Valley, above Tarras, 31 Dec 1949, McNeur (CHR); 

Cromwell Gorge, near railway station, H.H.Allan (CHR50088); Cromwell Gorge, E.Edgar (K); Alexandra, 

J.E.Holloway (OTA2964); Alexandra, A.Wall (CANTY1787). 

27. Luzula celata Edgar, New Zealand J. Bot. 4: 165 (1966) 

T: New Zealand, South Is., Canterbury, Rangitata Valley, near Potts River bridge, 29 Oct 1964, E.Edgar; holo: 

CHR149586; iso: K. 

Illustration: P.J.Lange, A.D.Wilton & C.Beard, New Zealand Bot. Soc. Newsl. 53: front cover (1998). 

Perennials, loosely tufted, forming loose cushions up to 3 cm tall; rhizome creeping to 

ascending, branched; stolons present, with base covered with membranous-scarious 

cataphylls with distinct veins. Leaves up to 4 cm long, c. 0.5–1.0 mm wide, involute to 

canaliculate; apex acute; margins ±smooth, usually densely ciliate. Stem abbreviated, much 

shorter than leaves, often not visible. Lower bract 1 (–3), usually 0.5–1.0 cm long, base and 

middle part herbaceous, otherwise ±membranous. Inflorescence reduced to a single (very 

rarely two) (3–) 6–16-flowered cluster. Bracteoles scarious, c. 1 mm long, ciliate. Tepals 

±equal, glabrous, ovate-lanceolate, 1.7–2.1 mm long, scarious with pale castaneous base and 

midrib. Stamens 3, equalling to slightly exceeding perianth; anthers 0.4–0.6 mm long; 

filaments 1.4–1.8 mm long. Capsule ±globular, subacute, pale brown, ±equalling perianth; 

capsule segments 1.8–2.0 × 1.7–1.8 mm. Seeds ellipsoidal, c. 1.0–1.2 mm long, c. 0.6–0.7 mm 

wide; appendage c. 0.1 mm long, fibrillate. 2n=12, fide E.Edgar, New Zealand J. Bot. 4: 165 

(1966). 

Endemic to South Island, New Zealand. 51: NZS. Sandy or shingly ground on old river 

terraces or in dune systems. Usually growing in, or near, cushions of Raoulia australis 

Hook.f. (Asteraceae). Map 53. 

61

62 


51. NEW ZEALAND SOUTH: South Is., Marlborough, Upper Awatere Valley, Muller Station, L.B.Moore 

(CHR77227); Ashburton R., Blowing Point, A.Wall (CANTY1737); Dunedin, Hoopers Inlet, L.B.Moore 

(CHR120151); Otago, Cromwell, Chafer Beetle Reserve, 210 m, A.D.Wilton 98223 (CHR, PRA); Otago, 

Crystall's Beach, Cooks Head, A.D.Wilton 98019 (CHR, PRA); Stewart Is. Mason Bay, E of Big Sandhill, 

H.Wilson (CHR). 

Known from about 15 sites in the South Island and one site in Stewart Is.; vulnerable. This 

species is extremely elusive, and is easily overlooked. 

28. Luzula pindica (Hausskn.) Chrtek & Křísa, Webbia 19: 6 (1964); Notes Roy. Bot. 

Gard. Edinburgh 25: 164 (1964) 

Luzula spicata β [unranked] pindica Hausskn., Mitt. Thür. Bot. Ver. 13–14: 33 (1899); Luzula bulgarica 

subsp. pindica (Hausskn.) Chrtek & Křísa, Bot. Not. 115: 308 (1962); Luzula spicata subsp. pindica 

(Hausskn.) Gamisans, Candollea 29: 42 (1974). T: Greece, Pindhos Mtns, Mt Zygos, Jul 1885, 

H.C.Haussknecht; syn: JE, W. 

Illustration: J.Chrtek & B.Křísa, Bot. Not. 115: 309, fig. 7 (1962). 

Maps: J.Chrtek & B.Křísa, Bot. Not. 115: 307, fig. 6 (1962); J.Chrtek & B.Křísa, Webbia 19: 8 (1964). 

Perennials (15–) 20–35 (–45) cm tall, densely caespitose, sparsely hairy; rhizome short, 

vertical; stolons absent. Basal leaves ±flat, c. 3.5–6 (–9) cm long, of variable width, some 

usually 3.5–5.0 mm wide; cauline leaves 2–4, remote, c. 2–5 cm long, with lower ones often 

3–4 mm wide and upper ones 2–3 mm wide, sometimes narrower (c. 1.0–1.5 mm wide). All 

leaves ±acute to subobtuse; margins densely papillose-serrulate. Lower bract sometimes 

remote, subherbaceous, very narrow, acute, c. 1–2 cm long, usually shorter than inflorescence; 

middle bracts conspicuous, membranous, whitish straw-brown, castaneous at base, 

lanceolate-aristate, usually 6–9 mm long. Inflorescence nodding (on arcuate upper part of 

stem), ±dense to inconspicuously interrupted at base, spike-like, usually 1.2–2.5 × c. 1 cm, of 

2–7 few-flowered clusters. Bracteoles scarious, often brownish at base, ±ciliate, ovatelanceolate, 

c. 2.5 mm long. Tepals subequal, glabrous, lanceolate, acuminate, usually 2.8–3.7 mm 

long, dark castaneous-brown, with narrow paler membranous margins. Stamens 6; anthers 

0.9–1.2 mm long; filaments 0.3–0.4 mm long; style 0.6–0.9 mm long; stigmas c. 1.1–1.7 mm 

long. Capsule ±ovoid, subacute, pale to dark castaneous-brown, ±exceeding perianth; capsule 

segments 2.5–3.0 × 1.9–2.2 mm. Seeds subglobular, pale to dark brown, 1.4–1.8 mm long, 

1.1–1.3 mm wide; appendage yellowish, c. 0.2 mm long, fibrillate. 2n=24, fide R.Franzén, in 

A.Strid & Kit Tan, Mount. Fl. Greece 2: 742 (1991). 

Southern Europe: Balkan Peninsula and S Italy. 13: ALB, BUL, GRC, ITA, YUC. Subalpine 

meadows, usually on silicic rocks, usually between 1800 and 2500 m. Map 54. 

13. ALBANIA: Permët, Biovizhdë, Nemerçka Mtns, A.Alston & N.Y.Sandwith 1811 (K). BULGARIA: Pirin 

Mtns, Banderica, below Mt Vikhren, c. 2200 m, J.Kirschner 4465 (PRA). GREECE: Macedonia, Samarina, 

Mt Smolikas, 2150 m, E.K.Balls & H.W.Gourlay 3418 (K); Thessalia, Mt Olympos, 2600–2650 m, A.Strid 

1505 (C, LD); Pindhos Mtns, Trikala, Dokimi, P.E.E.Sintenis [It. Thessal.] 844 (LD, PRC); Evritania, 

Karpenision, Mt Timfrestos, 2050–2100 m, L.-Å.Gustavsson 1626 (LD). ITALY: Basilicata, Mt Volturino, 

1830 m, O.Gavioli s.n. (FI). CRNA GORA: Javorje Planina [Mtns], Jul 1903, J.Rohlena (PRC); Štavna, Mt. 

Kom, Jul 1903, J.Rohlena (PR, LD). F.Y.R. MACEDONIA: Štrba, Šar Planina [Mtns], 2100 m, 18 Sep 

1922, K.Vandas (PR). 

At the sites with a common occurrence of L. pindica and L. spicata subsp. italica in the 

Balkans, intermediates between the two taxa are found. They are probably of a hybrid origin. 

A specimen from E Anatolia [Amasya, Mt Ak-dagh, J.F.N.Bornmüller [Pl. Exs. Anatol. Or.] 

884 (WU)] has most of the attributes of typical L. pindica but the material is insufficient for 

a sure identification. 

29. Luzula spicata (L.) DC., in J.B.A.P. de M. de Lamarck & A.P. de Candolle, Fl. 

Franç., 3rd edn, 3: 161 (1805) 

Juncus spicatus L., Sp. Pl. 330 (1753); Gymnodes spicata (L.) Fourr., Ann. Soc. Linn. Lyon, sér. 2, 17: 173 

(1869); Juncoides spicata (L.) Kuntze, Revis. Gen. Pl. 2: 725 (1891). T: Sweden, Lapland, C.Linnaeus 125; 

lecto: Flora Lapponica herbarium, Institut de France, Paris, designated here.


Luzula spicata (L.) DC. var. compacta E.Mey., Syn. Luzul. 19 (1823); Luzula compacta (E.Mey.) Dalla Torre 

& Sarnth., Farn- u. Blütenpfl. Tirol. 1: 437 (1906), non sensu orig., nom. superfl.; Luzula spicata f. compacta 

(E.Mey.) I.Grint., Fl. Rep. Soc. Roman. 11: 596 (1966). T: Bohemia, ‘Riesengebirge’ [the Krkonoše Mtns]; 

syn: A.J.Krocker, Fl. Siles. 5(1): tab. 52 (1787) [a mixture of elements belonging to subsp. spicata and subsp. 

conglomerata were given by E.Meyer; the name was validated through a reference to A.J.Krocker, Fl. Siles. 

5(1): 572 (1787)]. 

Luzula spicata var. relaxa Krylov, Fl. Alt. & Tomsk. Gub. 6: 1401 (1912). T: not indicated. 

Maps: E.Hultén, Kongl. Svenska Vetenskapsakad. Handl., ser. 5, 7(1): map 218 (1958); H.Meusel, E.Jäger & 

E.Weinert, Vergl. Chorol. Zentraleur. Fl. 1: 88 (1965). 

Perennials, usually (5–) 10–30 (–40) cm tall, densely caespitose; rhizome short; stolons 

absent. Basal leaves numerous, usually 1.0–2.5 (–3.5) mm wide; cauline leaves 2–3 (–4); 

basal leaves ±acute to obtuse (not swollen); cauline leaves and bract acute, with margins 

±densely papillose-serrulate. Lower bract subherbaceous or brownish membranous, shorter 

than or equalling inflorescence. Inflorescence nodding, rarely suberect, ±interrupted at base 

or not so, composed of clusters subtended by conspicuous middle bracts; middle bracts 

ovate-lanceolate, acuminate, membranous, ciliate. Tepals ±equal, glabrous, oblong-lanceolate, 

acuminate, brown, with indistinct paler margins distally. Stamens 6; style 0.2–0.4 mm long; 

stigmas 0.8–1.3 mm long. Capsules dark castaneous-brown, usually subacute. Seeds 

±ellipsoidal, ±paler brown, (0.8–) 0.9–1.1 (–1.3) mm long, (0.5–) 0.6–0.7 mm wide; seedcoat 

distinct, forming a pale apex; appendage whitish-yellowish, 0.1 (–0.2) mm long, 

fibrillate. 

A widely distributed complex of distantly disjunct populations confined to the Arctic and 

high altitudes in the N hemisphere mountains. Individual populations differ from one another 

in a series of mostly quantitative traits. The geographical variability is accompanied by an 

agmatoploid variation. The following 5 subspecies are recognized; some populations (in 

Turkey and the Alps) do not fit this pattern. The whole complex requires a detailed revision. 

A recent study (C.Garcia-Herran, in litt.) revealed a unique feature of the L. spicata group: 

partially fragmented ("aneuploid'') chromosome sets in populations or groups of populations. 

Note: Differences between the size of seeds, capsules and tepals given below and that 

reported in the literature is a consequence of a different method of measurement. Seeds (as 

everywhere in Luzula) are measured excluding the basal appendage, seen from above (dorsal 

view), tepals and capsule segments are first liberated from the inflorescence and measured 

separately. 

1 Anthers 0.4–0.5 (–0.7) mm long, ±equalling filaments 

2 Inflorescence usually interrupted at base 29a. subsp. spicata 

2: Inflorescence compact 

3 Seeds usually 0.9–1.2 mm long; capsule segments 1.5–2.1 mm long; 

inflorescence 1.2–1.5 × c. 1.0 cm 29b. subsp. conglomerata 

3: Seeds usually 0.7–0.9 (–1.0) mm long; capsule segments 1.2–1.7 mm 

long; inflorescence 0.7–1.3 × 0.7–0.8 cm 29c. subsp. mongolica 

1: Anthers (0.5–) 0.6–0.9 (–1.2) mm long, at least 1.5–2.0 times as long as 

filaments 

4 Capsule segments 2.0–2.8 mm long 29d. subsp. italica 

4: Capsule segments 1.4–1.9 mm long 29e. subsp. nevadensis 

29a. Luzula spicata (L.) DC. subsp. spicata 

Luzula obtusata Steud., Syn. Pl. Glumac. 2: 294 (1855). T: Greenland, ‘Friedrichthal Groenlandiae’, 

R.F.Hohenacker; syn: P, PR. 

Luzula spicata var. petraea Laest., Bot. Not. 1858: 145 (1858) [vel Nova Acta Regiae Soc. Sci. Upsal. 11: 

260–270 (1839), n.v.]. T: Njammats i Qvickjock, L.L.Laestadius; syn: UPS. 

Luzula spicata var. major Lange, Consp. Fl. Groenland. 128 (1880). T: [Greenland] ‘Disco i Lyngmarken, 

T. Fries; syn: n.v.; Tunugdliarfik-Fjord, A.Kornerup; syn: C. 

63

64 


Luzula spicata f. petraea Laest. ex Neuman, Sver. Fl. 665 (1901). T: Sweden [otherwise not indicated]. 

Luzula cusickii Gand., Bull. Soc. Bot. France 66: 295 (1919). T: USA, Oregon, Blue Mts., Anthony’s Ck, 

W.C.Cusick 2248; holo: LY, n.v. 

Luzula spicata var. nova Smiley, Univ. Calif. Publ. Bot. 9: 128 (1921); Luzula spicata subsp. saximontana 

Á.Löve & D.Löve, Univ. Colorado Stud. Ser. Biol. 17: 17 (1965). T: California. Sierra Nevada, Tulare Co., 

near Mineral King, F.V.Coville & F.Funston 1535; holo: n.v. 

Illustrations: J.Kirschner, Stud. Českoslov. Akad. Věd 1989(10): 12, fig. 1 (1989); J.Chrtek & B.Křísa, Webbia 

19: 2, fig. 1, Plate i, Feb (1964). 

Map: J.Chrtek & B.Křísa, Bot. Not. 115: 302, fig. 4 (1962). 

Plants usually 15–25 (–40) cm tall. Basal leaves usually up to 1.5–2.0 (–3.0) mm wide. 

Inflorescence lobate, ±interrupted at base or not so, often narrow, elongated, up to 3 × 1.5 cm. 

Tepals usually 2.2–2.8 (–3.3) mm long. Anthers (0.4–) 0.5–0.6 (–0.7) mm, ±equalling 

filaments; filaments c. 0.4–0.5 mm long. Capsule segments usually 1.6–2.3 × 1.0–1.2 mm. 

Seeds usually 0.9–1.1 × c. 0.6 (–0.7) mm; appendage 0.10–0.15 mm long. 2n=24 [24BL], 

fide H.Nordenskiöld, Hereditas 37: 334–335 (1951). 

Northern and Central Europe (the Alps, including southern Alps, the Krkonoše, the 

Pyrenees), N Great Britain, the N Urals region, Greenland, arctic and mountain regions of N 

America. 10: FIN, FOR, GRB, ICE, NOR, SVA, SWE. 11: AUT, CZE, GER, POL, SWI. 

12: FRA, SPA. 13: ITA, YUC-SL. 14: RUN. 30: WSB. 70: ASK, GNL, NWT, YUK. 

71: ABT, BRC, MAN. 72: LAB, NFL, QUE. 73: COL, IDA, MNT, ORE, WAS. 75: MAI, 

NWH, NWY, VER. 76: ARI, CAL, NEV. 77: NWM. Arctic and alpine tundra, on winderoded 

acidic soils. Map 55. 

10. GREAT BRITAIN: Easterness, Cairn Gorm, Coire an Lochan, A.Melderis 418 (S); SWEDEN: Torne 

Lappmark, Jukkasjärvi, Björkliden, 400 m, 22 Jul 1928, Levan (K). 11. CZECH REPUBLIC: Krkonoše 

Mtns, Mt Sněžka, Missbach [Petrak Fl. Bohem. Morav. Exs.] 138 (BRNU, PRC, PR). 12. FRANCE: Puyde-Dôme, 

Mont-Dore, Puy Ferrand, 1400–1800 m, P.Billiet [C.Magnier Fl. Select. Exs.] 988 (LD, PRC). 

14. RUSSIA NORTH: Archangelsk Region, Kanin Peninsula, mouth of Schoyna R., Sergienko [Gerb. Fl. SSSR] 

6105 (K). 70. GREENLAND: Fiskefjord, Qeqertat, J.Feilberg [Pl. Vasc. Groenl. Exs.] 689 (UPS); 

W Greenland, Neria, 61°33' N, 3 Jul 1931, J.Eugenius (LD, PR). 71. ALBERTA: Waterton Lakes Nat. Park, 

Mt Lineman, A.Breitung 14023 (S). 72. QUEBEC: E coast of Hudson Bay, Great Whale River, O.Hedberg 

2624 (UPS). 75. NEW HAMPSHIRE: Mt Lafayette, M.L.Fernald [Pl. Exs. Gray.] 191 (LD, PRC, W, WU). 

Variable. Some plants from Greenland are exceedingly robust (described as L. obtusata 

Steud.); the aberrant variation probably depends on edaphic conditions. Plants from W USA 

and SW Canada require further study; towards the south, the western plants show a tendency 

to broader leaves, longer tepals and broader seeds. The stamen and seed characters, however, 

are within the subsp. spicata limits. If the western plants prove to represent a distinct taxon, 

two names, var. nova Smiley, and subsp. saximontana Á.Löve & D.Löve, are referable to it. 

29b. Luzula spicata subsp. conglomerata (W.D.J.Koch) Murr, Magyar Bot. Lapok 

28(1929): 67 (1930) 

Luzula glomerata Miel. ex Huebener, Flora 22: 489 (1839); Luzula spicata var. conglomerata W.D.J.Koch, 

Syn. Fl. Germ. Helv., 2nd edn, 848 (1844) [‘L. conglomerata Mielichhofer’ mistakenly cited]. T: Austria, 

Salzburg, Gastein Tal, Rathausberg, M.Mielichhofer; lecto: PR, designated here; isolecto: W9004. 

Luzula spicata a. [unranked] subpediformis Schur, Enum. Pl. Transsilv. 683 (1866). T: [Romania] 'Auf der 

Keprereasze der Arpaser Alpen, am Keprereasze See', F.Schur; syn: n.v. 

Luzula spicata subsp. mutabilis Chrtek & Křísa, Bot. Not. 115: 302 (1962). T: Slovakia, Vysoké Tatry, 

Furkotský štít, K.Domin & V.Krajina [Fl. Čechosl. Exs.] 337; holo: PRC. 

Luzula spicata subsp. montsignatica P.Monts., Anales Inst. Bot. Cavanilles 21(2) [1963]: 478 (1964). 

T: [Spain] Montseny, Matagalls, 1650 m, 31 Jul 1949, O.Bolós; holo: BC113857. 

Luzula spicata f. minima [Schur ex] I.Grint., Fl. Rep. Soc. Roman. 11: 596 (1966), nom. inval. T: not 

designated. 

Illustrations: T.Szynal & J.Mądalski, Atlas Fl. Polsk. 2(2): 146 (1931); Chrtek & Křísa, Webbia 19: 2, fig. 2, 

Plate iii, iv (1964). 

Map: J.Chrtek & B.Křísa, Bot. Not. 115: 305, fig. 5 (1962).


Figure 14. Luzula spicata subsp. conglomerata. A, habit; B, partial inflorescence; 

C, perianth and capsule; D, capsule; E, seed; F, immature capsule with style and stigmas; 

G, H, tepals; I, bracteole; J, stem T.S.; K, leaf T.S. (A.Rehman s.n., KRA). Scale bars: 

A = 5 cm; B = 5 mm; C–I = 2 mm; J, K = 1 mm. Drawn by T.Szynal. Reproduced with 

permission, from T.Szynal & J.Mądalski, Atlas Fl. Pol. 2/2: 146 (1931). 

65

66 


Plants 7–15 (–20) cm tall. Basal leaves up to 1.5–2.0 mm long. Basal bract often ±equalling 

inflorescence. Inflorescence ±abbreviated, subglobose, not interrupted, rarely lobate, up to 

1.5 × 1.0 cm. Tepals usually 1.8–2.4 (–2.7) mm long. Anthers 0.3–0.5 (–0.6) mm long, 

±equalling filaments; filaments c. 0.4–0.5 mm long. Style 0.2 mm long; stigmas c. 1.0 mm 

long. Capsule segments 1.5–2.1 × c. 1.1–1.2 mm. Seeds 0.9–1.2 × c. 0.6 mm; appendage 

c. 0.1 mm long. 2n=12, fide H.Nordenskiöld, Hereditas 37: 334–335 (1951); J.Májovský & 

A.Murín, Karyotax. Prehl. Fl. Slov. 362 (1987); 2n=18 [6AL+12BL], 2n=24 [24BL], fide 

C.Garcia-Herran, in litt. Fig. 14. 

Mountains of C and SW Europe: the Carpathians, the Alps and N Italy, the Pyrenees and 

N Iberian mountains, Swiss Jura. 11: AUT, CZE, GER, POL, SWI. 12: FRA, SPA. 13: ITA, 

ROM, YUC-SL. 14: UKR. Oligotrophic, wind-eroded alpine tundra. Map 56. 

11. AUSTRIA: Kärnten, Hohe Tauern, Mallnitz, 2460 m, 19 Aug 1919, G.Beck (PRC); Kärnten, Heiligenblut, 

Pasterze Glacier, D.H.Hoppe (PR); Tirol, Sterzing, Riedberg, 2500–2700 m, R.Huter [Fl. Exs. Austro-Hung.] 

3914 (PR, PRC, UPS). POLAND: Tatry Wysokie, Rysy, 2200–2500 m, B.Pawłowski [Pl. Polon. Exs.] 84 

(K, KRA, KRAM). SLOVAKIA: Vysoké Tatry, Skalnaté pleso, 14 Jul 1943, F.Nábělek (SAV). 

SWITZERLAND: Valais, between Riffelberg and Gornergrat, 2500–3000 m, F.O.Wolf [A.Kneucker, Cyper. 

Junc. Exs.] 180 (PR). 13. ROMANIA: Bucegi Mtns, 2000 m, 5 Jul 1904, J.Prodan (PR); 25 Jul 1970, 

I.Morariu et al. (LD). 

Very variable. While in the E Alps subsp. conglomerata is easily distinguishable from subsp. 

spicata, in the W Alps the limit is not so clear-cut, and karyological variation has also been 

observed. The type of the name L. spicata subsp. montsignatica has anther length and 

anther/filament ratio at the upper limit of the subspecies; further study is needed. 

29c. Luzula spicata subsp. mongolica Novikov, Nov. Sist. Vyssh. Rast. 26: 34 (1989) 

T: Mongolia, Mongolian Altai, Munch-Chajrchan-Ula, 14 Aug 1978, Ogureeva; holo: MW; iso: LE. 

Luzula spicata var. kunawurensis D.Don, Trans. Linn. Soc. 18(3): 324 (1840). T: India, ‘Kunawur ad Lippa’ 

[probably: Li-phu-lie-khe-shan-khou, Uttarpradesh], J.F.Royle; holo: LIV, n.v.; iso: W. 

Illustrations: S.M.H.Jafri, Fl. Pakistan 138: 24, fig. 6 (1981); K.F.Wu, Fl. Reipubl. Popularis Sin. 13(3): 252, 

plate 61 (1997). 

Plants usually 8–13 (–20) cm tall. Basal leaves short, usually canaliculate, 1.0–1.5 (–2.8) mm 

wide. Lower bract brownish membranous, often remote. Inflorescence not interrupted, 

compact, subglobose, dark castaneous-brown, 7–13 × 7–8 mm. Tepals 1.9–2.4 (–2.7) mm 

long. Anthers 0.3–0.5 mm long; filaments 0.4–0.5 mm long; style c. 0.2–0.3 mm long; 

stigmas 0.8–1.1 mm long. Capsule segments 1.2–1.7 × 1.0–1.2 mm. Seeds 0.7–0.9 (–1.0) mm 

long, 0.5–0.6 mm wide; appendage 0.10–0.15 mm long, fibrillate. 2n=12, fide C.García 

Herran, in litt. 

Mountains of Middle Asia, the Himalayas, China (Tibet, Xinjiang, Sichuan, Yunnan) and 

Mongolia, the Altai and southernmost Siberia. 30: ALT, BRY, IRK, KRA, TVA, WSB. 

32: KAZ, KGZ, TZK, UZB. 34: AFG. 36: CHC, CHT, CHX. 37: MON. 40: JMK, PAK. 

Alpine tundra, scree, and margins of woodland patches near the timberline. Map 57. 

30. ALTAI: Kosh-Agatsh, South Tshuya Range, upper Elangash R., N.Frizen 1928 (PRA); SE Altai, South 

Tshuya Range, Tarkhata R., 14 Jul 1982, N.K.Kovtonyuk (PRA); Altai, Kuraiskiy Range, 2800 m, 30 Jun 1982, 

N.K.Kovtonyuk (PRA). 34. AFGHANISTAN: Wakhan, E of Tergan Qurum Valley, S of Bishutik Lake, 

4700 m, C.Grey-Wilson & T.F.Hewer 1530 (W). 36. CHINA SOUTH-CENTRAL: NW Sichuan, upper 

Yalong basin, Chola Shan, Dege to Garze, Maniganggo, 4850 m, G.Miehe, S.Miehe & U.Wündisch 94-421-1 (E). 

TIBET: Mekong – Nu Jiang [Salween] divide, NE of the pass above Bamda, 4800 m, G.Miehe & U.Wündisch 

94-41-19 (E). QINGHAI, Daban Shan, Nan Shan, P.Küpfer (NEU 95-492; 2n=12). SICHUAN: Xinjiang, 

Altaï, 2640 m, P.Küpfer (NEU 95-491; 2n=12). 37. MONGOLIA: Lake Khubsugul, Manku-Sardyk, 8 Aug 

1975, L.I.Malyshev & Tshepurnov (PRA). 40. INDIA: Kashmir, Mt Apharwat, 4200 m, R.R.Stewart 8690 

(S). PAKISTAN: Chitral, Barum Gol, South Barum Glacier, 4500 m, 3 Jul 1950, P.Wendelbo (S). 

Similar to subsp. conglomerata but seeds, capsule segments and inflorescence usually smaller.


29d. Luzula spicata subsp. italica (Parl.) Arcang., Comp. Fl. Ital. 713 (1882) 

Luzula italica Parl., Fl. Ital. 2: 309 (1857); Luzula spicata var. italica (Parl.) Rouy, Fl. France 13: 267 

(1912); Luzula spicata f. italica (Parl.) H.Lindb., Acta Soc. Sci. Fenn., n. s. B 1(2): 37 (1932). T: Corsica, 

Monte Renoso; syn: n.v.; Sardinia, Monte Genargentu, E.Requien; syn: FI; G.G.Moris; syn: n.v. 

Luzula bulgarica Chrtek & Křísa, Bot. Not. 115: 304 (1962); Luzula spicata subsp. bulgarica (Chrtek & 

Křísa) Gamisans, Candollea 29: 42 (1974). T: Bulgaria, Rila Planina, Edigeol, 2350 m, 1 Aug 1923, 

F.A.Novák; holo: PRC. 

Luzula stilbocarpa Kirschner & Křísa, Preslia 51: 336 (1979). T: Russia, Caucasus, Dombay, between 

Severnyi Priyjut and Lake Klukhorskoe ozero, 2000–2500 m, 14 Jul 1977, J.Kirschner, M.Šourková et al.; 

holo: PRC; iso: PRC. 

Illustrations: J.Kirschner & B.Křísa, Preslia 51: 337, fig. 2 (1979) as L. stilbocarpa; J.Chrtek & B.Křísa, 

Webbia 19: 5, fig. 3, Plate v (1964). 

Maps: J.Chrtek & B.Křísa, Bot. Not. 115: 307, fig. 6 (1962); Chrtek & Křísa, Webbia 19: 8 (1964). 

Plants usually 12–30 cm tall. Basal leaves usually canaliculate, up to 2.0 (–2.5) mm wide, 

usually densely ciliate at blade base. Inflorescence usually lobate and/or interrupted, c. 1.5–2.5 × 

1.0 cm. Tepals usually (2.0–) 2.3–2.8 (–3.0) mm long. Anthers (0.5–) 0.6–0.9 (–1.2) mm, 

usually 1.5–3.0 times as long as filaments; filaments 0.3–0.4 mm long; style 0.3–0.4 mm long; 

stigmas c. 1.0–1.5 mm long. Capsule segments usually (1.9–) 2.0–2.4 (–2.8) × 1.2–1.4 mm. 

Seeds (0.9–) 1.0–1.1 (–1.3) mm long, 0.6–0.8 mm wide; appendage c. 0.1 (–0.2) mm long, 

fibrillate. 2n=24 [24BL], fide J.Kirschner, ined.; 2n=16 [8AL+8BL], fide C.Garcia-Herran, 

in litt. 

Mountains of southern Europe from Corse, Sardinia and C Italy to Balkan Peninsula, Turkey, 

the Caucasus and NW Iran. 12: COR, SAR. 13: ALB, BUL, GRC, ITA, YUC. 33: NCS, TCS. 

34: IRN, TUR. Subalpine and alpine meadows, alpine tundra, usually above 2000 m. Map 58. 

12. CORSE: Mt D’Oro, cascades des Anglais, 1900 m, 16 Jun 1991, C.Garcia-Herran (NEU91-921; 2n=12). 

13. BULGARIA: Rodopi, Mt Musala, 19 Aug 1936, J.Dostál (PRC). CRNA GORA: Mt Maglić, Aug 1905, 

J.Rohlena (PRC). F.Y.R. MACEDONIA: Mt Luboten, 2500 m, 9 Sep 1922, K.Vandas (PR49527). 

BOSNIA-HERZEGOVINA: Herzegovina, Gacko, Mt Lebršnik, Aug 1913, K.Vandas (PR49524). ITALY: 

Abruzzi, Monti Sibillini, Vettore, 2200–2300 m, 10 Aug 1898, S.Sommier (FI). 33. RUSSIA: the Caucasus, 

Tyrnyauz, Mt Cheget, 2300–2500 m, 28 Jul 1981, V.Vašák (W). GEORGIA: Mt Tskhra-Tskharo, 2300 m, 

23 Aug 1991, P.Küpfer (NEU93-926; 2n=8AL+8BL). 34. TURKEY: Bursa, Uludag, 2 Aug 1944, 

M.Basarman (ISTF4571, LD); Kastambuli, Tossia, Giaurdag, 17 May 1892, P.E.E.Sintenis 

[It. Orient.] 3929 (LD, PR). 

A variable complex of regional populations characterized by long anthers, usually much 

longer than the filaments, usually interrupted or distinctly lobate inflorescence, and large 

capsule segments and seeds. Local populations often are relatively distinctive, deviating 

from the average values in a number of quantitative characters. However, numerous 

substantial overlaps of the quantitative character ranges do not allow recognition of the local 

types at a rank higher than variety. A unique feature is the multiple, fixed occurrence of 

2n=16=8AL+8BL in the Caucasus. 

29e. Luzula spicata subsp. nevadensis P.Monts., Anales Inst. Bot. Cavanilles 21(2): 

478 (1964) 

T: Spain, Sierra Nevada, Mulhacén, Chorreras Negras, 2800 m, 19 Jul 1923, P.Font Quer; holo: BC89906. 

Luzula hispanica Chrtek & Křísa, Novit. Bot. Delect. Seminum Horti Bot. Univ. Carol. Prag. 1965: 28 (1965). 

T: Spain, Sierra Nevada, Valle del Lanjarón, Peñon Elorrieta, 3200 m, 6 Aug 1930, Ceballos & C.Vicioso; 

holo: M19783. 

Plants usually 6–20 cm tall. Basal leaves flat or canaliculate, up to 2.0 (–2.5) mm wide. 

Inflorescence usually ±interrupted or at least lobate, up to 2.0 × 1.0 cm. Lower bract often 

equalling inflorescence. Tepals usually 1.8–2.4 mm long. Anthers (0.5–) 0.6–0.8 mm long, 

about 1.5–2.0 times as long as filaments; filaments 0.3–0.5 mm long; style 0.2–0.3 (–0.4) mm 

long; stigmas 1.0–1.3 mm long. Capsule segments usually 1.4–1.9 × c. 1.1–1.2 mm. Seeds 

0.8–1.0 mm long, 0.5–0.6 mm wide; appendage c. 0.1 mm long, fibrillate. 2n=24 [24BL], 

fide C.Garcia-Herran, in litt. 

67

68 


Mountains of Spain and the Great Atlas in N Africa. 12: SPA. 20: MOR. Alpine grasslands 

and open scree slopes. Map 59. 

12. SPAIN: Granada, Sierra Nevada, Picacho Veleta, 2800 m, E.Wängsjö & G.Wängsjö 3840 (LD); P.Porta & 

G.Rigo [It. iii Hisp.] 512 (WU); Sierra Nevada, Valle de Genil, Hoya de la Moya, 2600 m, E.Wängsjö & 

G.Wängsjö 3865 (LD); Madrid, Sierra de Gredos, Risco de la Cocina, E.Bourgeau [Pl. Esp.] 2555 (W); Sierra 

de la Demanda, Laguna Negra, 1930 m, 21 Jul 1990, C.Garcia-Herran (NEU91-1190; 2n=24); Huesca, 

Pirineos centrales, Formigal, 1700 m, 21 Aug 1992, C.Garcia-Herran (NEU92-892; 2n=24). 

20. MOROCCO: the Great Atlas, Jebel Toubkal, 3300–3600 m, P.H.Davis 55492 (E). 

A taxon very close to subsp. italica but with all floral parts smaller. The relationship of some 

inland Spanish populations and some Pyrenean plants remains unclear. 

30. Luzula pediformis (Chaix) DC., in J.B.A.P. de M. de Lamarck & A.P. de Candolle, 

Fl. Franç., 3rd edn, 3: 162 (1805) 

Juncus pediformis Chaix, in D.Villars, Hist. Pl. Dauphiné 1: 318 (ii 1786). T: France, ‘à Chaudun, à la 

Grangette’; syn: n.v.; ‘Dauphiny’, D.Villars; possible syn: LINN-SMITH; Col de la Coche, Lautaret, 

D.Villars; authentic material: GRM26858. 

Juncus nutans Chaix ex Vill., in J.-E.Gilibert, Syst. Pl. Eur. 1: 34 (1785), nom. inval., opus oppressus; Fl. 

Delphin. 34 (vi 1786); Luzula nutans (Vill.) Duval-Jouve, Bull. Soc. Bot. France 10: 80 (1863); Juncoides 

nutans (Vill.) Kuntze, Revis. Gen. Pl. 2: 724 (1891). T: [France] ’Challiol de Vieil, Embrun etc.’, D.Chaix; 

syn: ?GRM, n.v. 

Illustration: S.Pignatti, Fl. Ital. 3: 447 (1982). 

Perennials (25–) 30–55 (–85) cm tall, loosely caespitose. Rhizome long, horizontal to 

oblique, branched, c. 4–6 mm in diam.; upper part covered with dark castaneous-brown leaf 

sheath bases,; older sheath remains often forming dark brown fibrous cover; stolons absent. 

Stem thick, usually 1.5–2.3 mm in the middle, with upper part nodding. Basal leaves ±flat, 

usually 10–20 cm long, (4–) 5–6 (–8) mm wide; leaves of intravaginal shoots narrower; 

cauline leaves 3–4 (–5), usually 5–9 cm long; all leaves ±densely ciliate, densely papilloseserrulate, 

acute with hyaline apex. Lower bract usually narrow, herbaceous (rarely lacking), 

usually equalling or overtopping inflorescence. Inflorescence nodding, (15–) 20–45 (–80)flowered, 

spike-like, lobate, often interrupted at base, usually 2.0–3.5 (–5.0) × 1.0–1.5 cm, 

composed of 3–10-flowered clusters, with lower 1 (–2) clusters often remote, ±pedunculate. 

Bracteoles 2.5–3.5 mm long, ±ovate, acuminate, ciliate, brownish-membranous. Tepals 

subequal, castaneous brown with broad straw-brown membranous margins, c. 4.5–6.0 × 

1.5–2.0 mm; outer tepals long acuminate, entire to (later) denticulate; inner tepals usually 

denticulate near apex, mucronate. Stamens 6; anthers 2.2–2.9 mm long; filaments c. 0.5–0.7 mm 

long; style 1.8–2.2 mm long; stigmas usually 2.5–3.0 mm long. Capsule paler castaneousbrown, 

acute to ±acuminate, ±mucronate; capsule segments 3.7–4.8 × c. 2.5–2.7 mm. Seeds 

pale brown, broadly ovoid, 2.1–2.4 mm long (including a 0.2–0.4 mm long apical caruncle), 

c. 1.5 mm wide; basal appendage an indistinct paler patch c. 0.1 mm long; base fibrillate. 

2n=12, fide C.García-Herran, in litt.; C.Favarger & P.Küpfer, Collect. Bot. (Barcelona) 7: 

353 (1968). 

SW Europe, from SW Alps to the mountains of N, C and S Spain. 12: FRA, SPA. 13: ITA. 

Mountain grasslands. Map 60. 

12. FRANCE: Hautes Alpes, Lautaret, 14 Aug 1888, Pellat (UPS); Hautes Alpes, Montgenévre, Gondron, 

Lannes [C.Magnier, Fl. Sel. Exs.] 1314 (W); Alpi Graie, Col du Mont Cenis, J.Bonjean [H.G.Reichenbach, 

Fl. Germ. Exs.] 1314 (UPS, W); E Pyrenees, Cerdagne, Dorres, 1900 m, E.M.Sennen [Pl. Esp.] 2863 (W); 

Pyrenees, Gédre, Saugué, 1000 m, Bordère [K.G.Baenitz, Herb. Eur.] 2799 (PR, UPS). SPAIN: Aragon, 

Panticosa, M.Willkomm [It. Hisp. ii] 358 (K); Huesca, Port de Gavarnie, 2 Aug 1959, G.Een (S); Granada, 

Poblo, Sierra de la Sagra, 1900 m, E.Reverchon [A.Kneucker, Cyper. Junc. Exs.] 59 (PR, WU); Segovia, Sierra 

de Guadarrama, Cerro del Aguila, 1860 m, 17 Jul 1978, S.Rivas-Martínez (BCF46672); Asturias, Macizo, 

Occidental de los Picos de Europa, Vegarredonda, 1600 m, 28 Jul 1982, H.S.Nava (FCO13916). 13. ITALY: 

Piemont, Col della Maddalena, 30 Jun 1958, E.Müller (UPS); ‘montagnes du Piemont’, G.B.Balbis (LD).


31. Luzula caespitosa (E.Mey.) Steud., Syn. Pl. Glumac. 2: 294 (1855) 

Luzula pediformis var. caespitosa [Gay ex] E.Mey., Linnaea 22: 420 (1849); Juncoides caespitosa (E.Mey.) 

Kuntze, Revis. Gen. Pl. 2: 724 (1891); Luzula pediformis subsp. caespitosa (E.Mey.) Guinea, Fl. Santander 

355 (1953). T: Spain, Asturias, Cangas de Tineo, Pico de Arvas, 1835, M.C.Durieu [Pl. Sel. Hispan. Lusit.] 

216; lecto: K, designated here; isolecto: W. 

Luzula caespitosa subsp. sanabriae P.Monts., Anales Inst. Bot. Cavanilles 21(2) [1963]: 473 (1964). 

T: Spain, Zamora, Sierra Segundera, Rivadelago, Moncalvo, laguna de Lacillos, 1700–1800 m, Jun 1948, 

T.M.Losa & P.Montserrat; holo: BC114621; iso: BCF4132, 46673. 

Luzula caespitosa subsp. iberica P.Monts., Anales Inst. Bot. Cavanilles 21(2) [1963]: 473 (1964). T: Spain, 

Burgos, Cordillera Ibérica, Laguna Negra, sobre Neila, 1800 m, P.Font Quer 339; holo: BC63341. 

Map: P.Montserrat, Anales Inst. Bot. Cavanilles 21(2): map 4 (1964). 

Perennials usually 20–30 cm tall, loosely caespitose, densely hairy at base; rhizome long, 

±densely branched, subvertical to ascending, c. 2–3 mm in diam., often covered with greyish 

brown fibrous remains of old leaf sheaths. Stem up to 1 mm in diam., upper part nodding. 

Basal leaves usually ±canaliculate, usually 4–7 (–11) cm long, up to 2 mm wide, (leaves of 

new tillers and intravaginal shoots much narrower, c. 0.5 mm). Cauline leaves 2–3; blade 

usually 3–4 cm long, ±equalling or shorter than leaf sheaths. All leaves with ±papilloseserrulate 

margins; apex acute to acuminate. Lower bract herbaceous at base, ±brownish, 

1.0–2.0 (–2.5) cm long, usually exceeding inflorescence; middle bracts ovate-lanceolate, 

membranous, c. 6–9 mm long, ciliate. Inflorescence ±nodding, usually 4–9 (–13)-flowered, 

spike-like, usually congested, rarely with a ±remote basal cluster (or flower). Bracteoles 

ovate, membranous, ciliate, c. 2.0–2.5 mm long. Tepals ±equal, entire to sparsely denticulate 

at apex, usually 3.7–4.5 mm long, castaneous-brown with paler to ±membranous margins 

above. Stamens 6; anthers 2.0–2.9 mm long; filaments 0.3–0.4 mm long; style 1.8–2.5 mm 

long; stigmas 2.0–2.5 mm long. Capsule straw-brown to pale castaneous-brown, subglobose, 

subacute, ±mucronate; capsule segments 3.0–3.5 × 2.6–3.0 mm long. Seeds pale brown, 

broadly ovoid, usually 1.6–1.9 mm long (including a small paler apex c. 1.5 mm wide); basal 

appendage yellowish, c. 0.2 mm long, fibrillate. 2n=12 [12AL], fide C.Garcia-Herran, in litt. 

Mountains of NW and C Spain and EC Portugal. 12: POR, SPA. Rocky slopes and dry 

mountain grasslands, usually above 1600 m. Map 61. 

12. PORTUGAL: Beira Baixa, Serra de Estrela, Rua dos Mercadores, A.Fernandes, F.Sousa & J.Matos 4483 

(UPS); Serra de Estrela, Torre, Manteigas, 13–25 Jun 1987, A.Polatschek (W). SPAIN: Asturias, 

Leitariegos, Cueto de Arvas, P.Font Quer & W.Rothmaler [Fl. Iber. Sel.] 205 (BCF, S, UPS); Asturias, Pico 

de Arvas, Jul 1898, M.Gandoger (W); Palencia, Pico de Curavacas, 16 Jul 1990, C.Garcia-Herran (NEU90- 

1038; 2n=12); León, Picos del Mampodre, Cervunal de Valverde, 1800–1900 m, 5 Aug 1952, T.M.Losa & 

P.Montserrat (BCF4129); León, Hayedo del Gildar, 1900 m, Garcia-Gonzales (JACA109685); 

Madrid/Segovia, Sierra de Guadarrama, Siete Picos, Cercedilla – La Granja, 2080–2120 m, V.J.Aran & 

M.J.Toha (JACA172898, MA); Soria, Laguna Negra, 21 Jul 1991, C.Garcia-Herran (NEU91-1089; 2n=12); 

Soria, Coraleda, Pico de Urbión, 2000–2200 m, J.L.Benito (JACA19895). 

A species variable in the size of its vegetative parts and in the shape and size of its 

inflorescence. The variation shows considerable overlaps between local populations, and 

extremes are scattered throughout the species’ range. Thus, the aberrant plants described by 

P.Montserrat as subspecies are here subsumed within the species. 

32. Luzula ecuadoriensis Balslev, Fl. Ecuador 11: 11 (1979) 

T: Ecuador, Cotopaxi, 3550 m, 1939, E.Asplund 6494; holo: S; iso: S. 

Illustration: H.Balslev, Fl. Neotrop. Monogr. 68: 62, fig. 31 (1996). 


Perennials 50–75 cm tall, sparsely hairy to subglabrous; rhizome ascending. Basal leaves 

(5–) 10–20 (–25) cm long, usually 6–10 mm wide; cauline leaves usually 3–5, with sheath to 

7 cm long, usually 9–15 cm long, up to 10 mm wide; all leaves acute with densely papilloseserrulate 

margins. Lower bracts similar to leaves, up to 17 cm long, distally gradually shorter 

(lower inflorescence branches often in axils of upper or middle cauline leaves). Inflorescence 

an interrupted ±drooping panicle in upper 1/3 (– 1/2) of plant height, composed of dense 

oblong-ovoid partial inflorescences on long primary peduncles; further composed of 

69

70 


cylindrical to oblong spike-like clusters 1–2 × 0.4–0.8 cm. Bracteoles membranous, 

±lanceolate, 1.5–2.0 mm long, fimbriate-ciliate above. Tepals ±castaneous brown, subequal, 

with outer tepals (2.3–) 2.5–3.0 mm long, lanceolate, acuminate, glabrous or very sparsely 

ciliate; margin pale, indistinct to ±absent. Stamens usually 3, sometimes 4–5; anthers 0.4–0.5 mm 

long; filaments 0.6–0.7 mm long; style 0.4–0.5 mm long; stigmas c. 1.0–1.2 mm long. 

Capsule subglobose to obovoid, paler castaneous brown; capsule segments 1.5–1.8 × c. 1.2 mm, 

glossy. Seeds c. 1.0 mm long, 0.5–0.6 mm wide; seed-coat distinct, forming a pale apex; 

basal appendage c. 0.1 mm long, fibrillate. Fig. 15. 

Endemic to Ecuador and Peru. 83: ECU, PER. Páramos and subpáramos or rocky slopes 

between 3300 and 3900 m. Map 62. 

83. ECUADOR: Azuay, Cuenca, Angas, 3400 m, H.Balslev 1509 (NY); Pichincha, Mt Pichincha above Quito, 

3700 m, H.Balslev 23615 (AAU, NY). PERU: Cajamarca, 2700–3700 m, B.Becker & F.M.Terrones 1225 

(AAU); Lambayeque, Incahuasi, Cerro Negro, 3300 m, A.Sagástegui et al. 12827 (AAU, F, MO). 

33. Luzula excelsa Buchenau, Abh. Naturwiss. Vereine Bremen 4: 126, tab. 4, fig. 1–8 

(1874) 

Juncoides excelsa (Buchenau) Kuntze, Revis. Gen. Pl. 2: 724 (1891). T: Bolivia, La Paz, Lancha de 

Cochipata, 2700–3000 m, 1860, G.Mandon 1449; lecto: GOET, fide H.Balslev, Fl. Neotrop. Monogr. 68: 61 

(1996); isolecto: BM, BR, F, GH, K, P, PR, S, US. 

Luzula hieronymi Buchenau & Griseb., in A.H.R.Grisebach, Symb. Fl. Argent. 318 (1879); Juncoides 

hieronymi (Buchenau) Kuntze, Revis. Gen. Pl. 2: 734 (1891). T: Argentina, Tucumán, Cienega, 1874, 

P.G.Lorentz & G.Hieronymus 735; lecto: GOET, fide H.Balslev, Fl. Neotrop. Monogr. 68: 61 (1996); isolecto: 

CORD, NY, K, S. 

Luzula hieronymi Buchenau & Griseb. var. pusilla Castillon, Bol. Mus. Hist. Nat. 7: 19 (1926). T: Argentina, 

Tucumán, Tafí, Cumbres Calchaquies, 4200 m, 29 Jan 1907, M.Lillo 5600; lecto: LIL, fide H.Balslev, Fl. 

Neotrop. Monogr. 68: 61 (1996); photo: PRA. 

Luzula hieronymi Buchenau & Griseb. f. bonariensis Barros, Darwiniana 10: 335 (1953). T: Argentina, 

Buenos Aires, Pigüé, Sierra de Curumalan, A.Burkart 4875; lecto: SI, fide H.Balslev, Fl. Neotrop. Monogr. 

68: 61 (1996). 

Luzula castellanosii Barros, Darwiniana 10: 349 (1953). T: Argentina, Salta, Cafayate, Sierra de los 

Quilmes, 26 Jan 1943, A.Castellanos; holo: BA46656. 

Illustrations: F.G.P.Buchenau, Abh. Naturwiss. Vereine Bremen 4: plate 4 (1874); F.G.P.Buchenau, in 

H.G.A.Engler, Pflanzenr. (iv.36) 25: 76, fig. 49 (1906); M.Barros, Darwiniana 10: 334, fig. 16, 337, fig. 17 

(1953); H.Balslev, Fl. Neotrop. Monogr. 68: 64, fig 33(1996). 

Perennials 40–100 cm tall, hairy to sparsely hairy; rhizome ascending to ±erect, rarely 

±creeping. Basal leaves usually up to 25 cm long, (5–) 8–13 (–20) mm wide; cauline leaves 

usually 3–6, with upper ones to 17–20 cm long and 13 mm wide; all leaves acute, with 

densely papillose-serrulate margin. Primary bracts long, leaf-like, herbaceous, often ±equalling 

their partial inflorescences (lower inflorescence branches often in axils of upper cauline 

leaves). Inflorescence a loosely branched drooping interrupted panicle constituting 1/3– 1/2 of 

plant height, composed of long pedunculate interrupted partial panicles of numerous small, 

cylindrical clusters usually 5–8 (–15) × 3–4 (–5) mm. Bracteoles membranous, ovate, 

acuminate, c. 1 mm long, fimbriate-ciliate above. Tepals straw-brown to pale castaneous brown, 

(1.3–) 1.5–1.8 (–2.2) mm long, ±equal, ±glabrous, with a distinct membranous or paler 

margin. Stamens 3; anthers 0.3–0.5 mm long; filaments 0.5–0.7 mm long; style 0.2–0.4 mm 

long; stigmas 0.5–0.7 mm long. Capsule ±ovoid, ±trigonous, acute, straw-brown to brown; 

capsule segments 1.4–1.6 × 0.9–1.1 mm. Seeds ellipsoidal, distinctly dark apicate, 0.8–0.9 mm 

long, 0.5–0.6 mm wide; appendage whitish, 0.10–0.15 mm long, fibrillate. Fig. 16. 

From southern Peru and Bolivia to Argentina and northern Patagonia. 83: BOL, PER. 85: 

AGE, AGW. Open mountain rocky slopes and open woodlands, 1400–3500 m. Map 63. 

83. BOLIVIA: Noryungas, Unduari, 3100–4000 m, O.Buchtien 2601 (GH, M, NY, UPS, US); La Paz, Murillo, 

Pongo, 3600 m, E.Asplund 1049 (UPS). 85. ARGENTINA NORTHEAST: Buenos Aires, Sierra de 

Curamalan, A.Fabris & H.Schwabe 4909 (CTES). ARGENTINA NORTHWEST: Catamarca, El Candado, 

P.Joergensen 1350 (SI); Tucumán, Chicligasta, Las Pavas, 3200 m, S.Venturi 3053 (S, US); Tucumán, Tafí del 

Valle, 2300 m, K.Hueck 15 (SI); Jujuy, Capital, Quebrada de Yala, A.L.Cabrera 27379 (SI); Catamarca, 

Andalgala, Alto Blanco, P.Joergensen 1564 (LIL); La Rioja, La Vega de La Hoyada, 2700-2800 m, 22 Jan


Figure 15. Luzula ecuadoriensis. General habit. (E.Asplund 6494, S). Scale bar: 5 cm. 

Drawn by K.Tind. Originally published in G.Harling & B.Sparre (eds.), Fl. Ecuador 11: 12, 

fig. 3 (1979), reproduced with permission, from H.Balslev, Fl. Neotrop. Monogr. 68: 62, 

fig. 31 (1996), © Lunds Botaniska Förening, © 1996, The New York Botanical Garden. 

71

72 


Figure 16. Luzula excelsa. A, habit; B, terminal part of partial inflorescence. 

(E.W.D.Holway & M.M.Holway 619, US). Scale bars: A = 5 cm; B = 2 mm. Drawn by 

K.Tind. Reproduced with permission, from H.Balslev, Fl. Neotrop. Monogr. 68: 64, fig. 33 

(1996), © 1996, The New York Botanical Garden.


1908, F.Kurtz 15129 (CORD). ARGENTINA SOUTH : Chubut. Tehuelches, Lago Vintter, 29 Jan 1995, 

E.Nicora 10083 (SI); Neuquén, Ñorquín, Copahué, 2000 m, C.A.O’Donnell 110458 (SI). 

A variable species with its highest diversity in N Argentina. Luzula castellanosii seems to be 

a depauperate, narrow-leaved form of L. excelsa. 

34. Luzula mendocina Barros, Darwiniana 10: 322 (1953) 

T: Argentina, Mendoza, Tunuyán, Cerro de las Piedras, 2220 m, 23 Mar 1935, A.Ruíz Leal 3105; lecto: 

LIL46294A, designated here; isolecto: LIL46294B. 


Perennials 25–60 cm tall, sparsely hairy caespitose; rhizome short; stolons absent. Basal 

leaves usually 12–15 cm long, 3.5–6.5 mm wide. Cauline leaves 2–3; upper ones usually 

short, 3–6 cm long, 2.5–3.5 mm wide. All leaves acute-subulate; margins with sparse reddish 

papillae. Primary bracts herbaceous, of variable length, usually 2.5–5.0 cm long. 

Inflorescence panicle composed of remotely interrupted, relatively dense, long pedunculate 

secondary panicles. Secondary panicles 2–4 × 1–2 cm, with lowermost often 10–15 cm below 

terminal one, of 3–10 spike-like dense clusters. Bracteoles scarious-membranous, 1–2 mm 

long, ±lanceolate, not or sparsely ciliate. Tepals subequal, entire, glabrous, lanceolate, 

acuminate; inner tepals 2.0–2.5 mm long; outer tepals 2.1–2.8 mm long. Stamens 6, rarely 

1 or 2 rudimentary; anthers 0.6–0.8 mm long; filaments 0.3–0.4 mm long; style 0.3–0.4 mm 

long; stigmas 0.8–1.1 mm long. Capsules narrowly ovoid, ±acute, brown, shining, shorter 

than perianth; capsule segments 1.5–1.7 × 1.0–1.1 mm. Seeds oblong-ovoid, c. 0.8–1.0 mm 

long, 0.6–0.7 mm wide; appendage indistinct, base fibrillate. 

Central Argentina (Mendoza, Cordoba, San Luis and Buenos Aires provinces). 85: AGE, 

AGW. Mountain grasslands, 700-3000 m. Map 64. 

85. ARGENTINA NORTHEAST: Buenos Aires, Sierra de la Ventana, Partido de Tornquist, Cerro de la 

Nerilana, 24 Mar 1948, A.L.Cabrera 54 (SI). ARGENTINA NORTHWEST: Mendoza, Tunuyán, Cuchilla 

del Cerro Bayo, 3000 m, A.Ruíz Leal 3085 (para: LIL); Tunuyán, Rincón Colorado, 3000 m, A.Ruíz Leal 1279 

(para: LIL46292A, B); Tunuyán, Las Cuevas, A.Ruíz Leal 3195 (LIL); Tunuyán, Bella Vista, 2300 m, G.Covas 

(SI18003); San Luís, La Carolina, A.Burkart 10777 (SI); Córdoba, Pampa de Achala, 13–14 Dec 1945, 

A.T.Hunziker 6425 (CORD). 

Distinct in having several, long pedunculate, ±dense secondary panicles ('heads'); it is unique 

in this respect among hexandrous taxa. 

35. Luzula masafuerana Skottsb., Nat. Hist. Juan Fernandez 2: 775 (1953) 

Luzula racemosa subsp. insularis Skottsb., in C.J.F.Skottsberg, Nat. Hist. Juan Fernandez 2: 110 (1922); 

Luzula insularis (Skottsb.) Skottsb., Veröff. Geobot. Inst. Rübel Zürich 3: 89 (1925), nom. illeg., non (Briq.) 

Prain (1921). T: Juan Fernández, Masafuera, Las Torres, 1100–1350 m, C.J.F.Skottsberg & I.Skottsberg 

408; lecto: S, designated here; isolecto: BM, K, LD, UPS. 

Illustration: C.J.F.Skottsberg, in C.J.F.Skottsberg, Nat. Hist. Juan Fernandez. 2: 110, fig. 5, b-e (1922). 

Perennials up to 35–40 cm tall, robust, densely hairy caespitose; rhizome short; stolons 

absent. Basal leaves usually 4–7 cm long, 4–9 mm wide, flat; cauline leaves 2 (–3), usually 

(3–) 5–7 cm long and 3.5–5.0 mm wide; all leaves acute, with papillose-serrulate margins. 

Lower bracts herbaceous, usually 2.5–6.0 cm long, exceeding inflorescence. Inflorescence 

erect to slightly bent, congested, compact, lobulate, or subdivided in 2–3 ovoid clusters and 

±elongated, 2.0–3.5 (–4.0) cm long. Bracteoles ±ovate, membranous, 1.0–1.5 mm long, 

densely fimbriate-ciliate. Tepals ±equal, lanceolate, distinctly acuminate, 2.2–3.0 mm long, 

±brown, ±glabrous to very sparsely ciliate. Stamens 3; anthers 0.3–0.6 mm long; filaments 

0.5–0.6 mm long; style c. 0.3 mm long; stigmas 1.0–1.2 mm long. Capsule ±ovoid, subacute, 

castaneous brown; capsule segments 1.7–2.0 × c. 1.3 mm. Seeds ellipsoidal, brown, 1.0–1.1 mm 

long, 0.6–0.7 mm wide; appendage absent; base fibrillate. 

Endemic to the Juan Fernández Is. 85: JNF. Found on rocks and rocky slopes. Map 65. 

85. JUAN FERNÁNDEZ IS.: Masafuera, N slope of Quebrada Casas, 900 m, 17 Jan 1986, T.Stuessy et al. 

9048 (OS); Quebrada de las Casas, ladera mirando al Sur Riscos, Muñoz & E.Sierra 7400 (OS); Masafuera, 

Quebrata Pasto, 1060–1100 m, 29 Jan 1986, T.Stuessy 9411 & T.Lammas (OS); Masafuera, Las Torres, 

73

74 


1370 m, 29 Jan 1955, I.Skottsberg & C.J.F.Skottsberg 399 (S); Masafuera, Cuchara, above Correspondencia 

Camp, 1350 m, I.Skottsberg & C.J.F.Skottsberg 162 (S); Masafuera, Quebrada Guaton, valley toward Cordon 

Atravesado, 1230 m, A.Landero & L.Gaete 9108 (OS); Masafuera, 1200–1300 m, C.J.F.Skottsberg 28 (UPS). 

36. Luzula alopecurus Desv., J. Bot. (Desvaux) 1: 159 (1808) 

Juncoides alopecurus (Desv.) Kuntze, Revis. Gen. Pl. 2: 734 (1891). T: ‘ad fretum magellanicum’; 'Magell.', 

P.Commerson; syn: P. 

Luzula villosa Wikstr., Kongl. Vetensk. Akad. Handl. 1823(2): 276 (1823). T: ‘Terra del Fuego et Falklands 

Islands’; syn: not located in S or UPS. 


Darwiniana 10: 339, fig. 18 (1953). 


Perennials usually 10–35 cm tall, densely hairy caespitose; base often purplish, robust; 

rhizome short; stolons absent. Stem rigidly erect, (0.6–) 0.8–1.2 mm in diam. below 

inflorescenceBasal leaves densely ciliate, numerous, 8–15 (–18) cm long, 4–5 (–6) mm wide; 

cauline leaves unevenly distributed, usually with one near stem base and 1 or 2 close to 

inflorescence, 6–10 cm long, often overtopping inflorescence; all leaves acute, with margins 

papillose-serrulate. Lower bract herbaceous, c. 2–5 cm long, usually overtopping inflorescence; 

upper bracts scarious-brownish, fimbriate-ciliate, up to 2 cm long. Inflorescence erect, 

densely congested, c. 1.5–2.5 cm long, of whitish villose appearance when younger, rarely 

with a small separate flower cluster at base. Bracteoles inserted c. 0.5–1.0 mm below flower; 

upper bracteoles narrowly lanceolate with a filiform tip, c. 3.0–3.5 mm long (often as long as 

flowers), densely fimbriate–ciliate. Tepals subentire, with outer ones densely and inner ones 

usually very sparsely long ciliate, narrowly lanceolate, acuminate with filiform tip, ±unequal 

(inner tepals 2.5–2.9 mm long, outer tepals 2.9–3.7 mm long), purplish castaneous brown to 

straw brown at base, with broad membranous border. Stamens 6; anthers 0.4–0.7 mm long; 

filaments 0.5–0.9 mm long, longer than anthers; style 0.1–0.2 mm long; stigmas c. 1.0 mm 

long. Capsule oblong-ovoid, ±trigonous above, ±acute, castaneous brown, shorter than 

perianth; capsule segments 1.8–2.0 × 1.3–1.4 mm. Seeds ellipsoidal, with distinct yellowish 

seed-coat and yellowish apex, dark castaneous, 0.9–1.0 mm long, 0.5–0.6 mm wide; 

appendage yellowish, 0.1–0.2 mm, fibrillate. 2n=24, fide D.M.Moore, Fl. Tierra del Fuego 

325 (1993). 

Endemic to southernmost Patagonia, Tierra del Fuego and the Falkland Islands. 85: AGS, 

CLS. 90: FAL. Grasslands and rocky slopes. Map 66. 

85. ARGENTINA SOUTH: Santa Cruz, C.Burmeister 173 (SI); Tierra del Fuego, Ushuaia, Río Piper, 

R.Santesson 411 (S); Isla Soledad, Seal Rookery, E.A.Ulibarri, M.Dimitri & E.Orfila 1039 (SI). CHILE 

SOUTH: Magallanes, Punta Arenas, Loretto, 9 Dec 1903, Scott Elliott (BM); Tierra del Fuego, Porvenir, 

R.Santesson 1565 (S); Tierra del Fuego, Sierra Carmen Silva, N side of Río Chico valley, D.M.Moore & 

R.N.Goodall 225 (H, RNG, UPS). 90. FALKLAND IS.: West Falkland Is., Roy Cove Ck, Vallentin 84 (K); 

Port Stanley, C.J.F.Skottsberg 90 (BM); 4 Jan 1908, C.J.F.Skottsberg 28 (S); East Falklands, Kidney Is., 

Sladen 5/49 (BM). 

37. Luzula brachyphylla Phil., Linnaea 33: 268 (1865) 

Luzula racemosa f. brachyphylla (Phil.) Fuentes, Bol. Mus. Nac. Hist. Nat. 90: 137 (1917). T: Chile, Chiloë 

Is., Chacao, Fonck 44; holo: SGO. 

Luzula patagonica Speg., Rev. Fac. Agron. Univ. Nac. La Plata 3: 577 (1897); Juncoides patagonica (Speg.) 

Kuntze, Revis. Gen. Pl. 2: 734 (1891). T: Argentina, Santa Cruz, Lago Argentino, Arroyo Leona, 

C.Spegazzini; syn: S. 

Luzula chilensis f. frequentior Barros, Darwiniana 10: 321 (1953). T: Argentina, Chubut, valle Lonquíñeu, 

Los Menucos, 8 Feb 1945, A.Castellanos; lecto: LIL114298, designated here. 

Luzula correae Barros, in M.N.Correa, Fl. Patagon. 2: 118 (1969). T: Argentina, Santa Cruz, Calafate, 

M.N.Correa 2962, L.Mendonza & C.Movia; holo: BAB, photo PRA. 

Perennials 15–30 (–55) cmtall, loosely caespitose, sparsely hairy; rhizome short or ascending; 

stolons absent. Basal leaves 4–8 (–17) cm long, 3–4 mm wide; cauline leaves 1 or 2, usually 

3–7 cm long; all leaves ±acute, with papillose-serrulate margins. Bracts erect to erecto-


patent, 1.5–2.5 (–4.0) cm long, equalling or overtopping their partial inflorescence, with 

lower bracts herbaceous and upper ones scarious. Inflorescence erect, rarely slightly 

nodding, interrupted, 2.5–6 cm long, of 3–6 (–10) clusters, with lower clusters pedunculate 

and upper ones subsessile; all clusters elongated, narrow (c. 4 mm in diam.), cylindrical 

composed of subclusters, many-flowered. Bracteoles c. 1 mm long, ovate, sparsely ciliate, 

scarious. Tepals ±lanceolate, subequal, glabrous; outer tepals acuminate, subaristate; inner 

tepals ±mucronate, 1.7–2.0 (–2.3) mm long; base castaneous; margin paler or membranous. 

Stamens 3 or 4–6; anthers 0.5–0.8 mm long; filaments 0.3–0.4 mm long; style 0.1–0.2 mm 

long; stigmas c. 1 mm long. Capsule ±ovoid, ±acute, deep brown; capsule segments 1.6–1.8 × 

1.0–1.1 mm long. Seeds dark, ellipsoidal, 0.9–1.0 (–1.1) mm long, 0.6–0.7 mm wide; seedcoat 

not distinct; apex not distinct; appendage c. 0.05 mm long, fibrillate. Fig. 17. 

Southern Chile and Southern Argentina. 85: AGS, CLS. In grasslands, usually at lower 

altitudes. Map 67. 

85. ARGENTINA SOUTH: Tierra del Fuego, Río Azopardo, P.Dusén 584 (UPS); Laguna Patos near Río 

Carbon, 4 Jan 1909, C.J.F.Skottsberg 943 (S, UPS); Chubut, Trevelin, T.Meyer 9277 (LIL); Chubut, José de 

San Martín, 530 m, 22 Jan 1902, Hözberg (S); Neuquén, Ñorquín, Copahué, 2000 m, C.A.O’Donnell 2097 

(LIL); Neuquén, Pino Hachado, A.Burkart 9651 (SI). CHILE SOUTH: Magallanes, Cerro Paine, S.F.Anilot 

6094 (SGO); Aysén, Laguna San Rafael, 3–4 m, C.Muñoz 6147 (SGO); Los Lagos, Mechuque, F.W.Pennel 

12622 (SGO); Los Lagos, San Rafael, 1921, C.M.Hicken (SI 10974). 

Most plants from Chile and from the southern part of the range are triandrous, with 

occasional occurrence of flowers with 4–6 stamens. Plants from Santa Cruz, Chubut and 

Neuquén, Argentina, are predominantly hexandrous (flowers with 3–5 stamens sometimes 

found in the same inflorescence as hexandrous ones). Differences in stamen number is the 

only significant variation found within the species (the names L. patagonica and L. correae 

refer to the predominantly hexandrous form). 

38. Luzula chilensis Nees & Meyen ex Kunth, Enum. Pl. 3: 312 (1841) 

Juncoides chilensis (Kunth) Kuntze, Revis. Gen. Pl. 2: 734 (1891). T: Chile, ‘ad Maipú fluv., alt. 10000 

ped.’, 1831, Meyen; syn: B, destroyed, see F.G.P.Buchenau, Abh. Naturwiss. Vereine Bremen 6: 418, Plate iv, 

fig. 1, a–d (1879) [et Junc. S.-Amer., 1879]; Chile, Bíobío, Antuco, E.F.Poeppig [Coll. Pl. Chil. III] 105; syn: 

BM, P, S. 

Luzula rigida Phil., Linnaea 33: 267 (1865); Luzula chilensis f. rigida (Phil.) Fuentes, Bol. Mus. Nac. Hist. 

Nat. 90: 138 (1917). T: Chile, Colchagua, ‘in montibus subalpinis Talcaregué, Feb 1831, [J.Gay] 284; 

holo: SGO. 

Luzula chilensis f. longifolia Barros, Darwiniana 10: 322 (1953). T: Argentina, Neuquén, Río Alumine, 

Otto Asp. 98; syn: BAF; Chubut, Río Percey, 6 Feb 1945, A.Castellanos; syn: LIL114302. 

Perennials, usually 12–25 (–55) cm tall, caepitose, sparsely to ±densely hairy; rhizome short; 

stolons absent. Basal leaves (5–) 6–8 (–12) cm long, 2.5–3.5 (–6) mm wide; cauline leaves 2, 

usually short, 1.5–3.0 cm long; all leaves acute, with margins distinctly papillose-serrulate. 

Lower bract membranous-brownish, rarely herbaceous, usually 1.5–2.0 cm long, ciliate, 

±equalling inflorescence. Inflorescence usually congested as a single slightly nodding or 

±suberect head c. 2 × 1.5 cm, of 5–6 clusters (usually 3–6-flowered); less often clusters 

distinct, ±interrupted, inflorescence about 5 cm long. Bracteoles scarious with straw-brown 

base, ovate, acuminate, fimbriate-ciliate, 1.5–2.0 mm long. Tepals ±equal, glabrous, 

lanceolate, c. 2.7–3.3 mm long, acuminate-aristate, pale castaneous brown below and on 

midrib, otherwise membranous to pale straw-brown. Stamens 6; anthers usually 0.6–0.9 mm 

long; filaments 0.4–0.5 mm long; style 0.3–0.4 mm long; stigmas 1.2–1.6 mm long. Capsule 

ovoid to subglobular, ±obtuse, castaneous brown, shining, equalling or shorter than perianth; 

capsule segments 2.4–3.0 × 1.5–2.1 mm. Seeds paler castaneous brown; seed-coat distinct, 

forming a yellowish apex, 1.2–1.5 mm long, (0.9–) 1.0–1.2 mm wide; appendage (0.1–) 0.2 mm 

long, fibrillate. Fig. 17. 

Most common in S and C Chile and S and W Argentina, reaching Valparaiso (district of 

Aconcagua) in C Chile. 85: AGS, AGW, CLC, CLS. Distribution imperfectly known. Wet 

mountain slopes and grasslands, usually below 2000 m, rarely above 3000 m. Map 68. 

75

76 


Figure 17. Luzula chilensis. A, habit; B, flower; C, tepal and stamen; D, seeds. 

L. brachyphylla. E, habit; F, perianth with unripe capsule; G, tepal and stamen; H, leaf apex 

and indumentum. (A–D, E.Wall 1727, S; F.W.Pennel 12442, S, SGO; E–H, C.J.F.Skottsberg 

943, S, UPS; A.Burkart 9651, SI). Scale bars: A, E = 5 cm; C, G = 1.5 mm; B, F = 2 mm; 

D = 1 mm; H = 1 mm. Drawn by © E.Smrčinová.


85. ARGENTINA NORTHWEST: Mendoza, Malarhue, Valle de Las Leñas, R.Kiesling & A.Donath 7262, 

25 Dec 1989 (SI, MCNS). ARGENTINA SOUTH: Neuquén, Cordillera del Viento, Cerro las Yeguas, 

A.Pastore 46 (S); Chubut, Lago Vintter, 1360–1450 m, S.Halloy A-93 (LIL); Santa Cruz, Río de las Vueltas, 

Estancia Pérez, 1200 m, H.Sleumer 1382 (LIL); Santa Cruz, Lago Argentino, 400 m, P.Dusén 5777 (S, SI, UPS). 

CHILE CENTRAL: Valparaíso, Petorca, Río Sobrante, 3100–3300 m, J.L.Morrison 17310 (S, SI); La 

Araucania, Cautín, Volcán Llaima, 1200–1300 m, E.Werdermann 1232 (LIL, S, SI); Valparaíso, Aconcagua, 

Portillo, 3000 m, E.Wall 1727 (S). CHILE SOUTH: Ñuble, Baños de Chillán, 2100 m, F.W.Pennel 12442 

(S, SGO); Libertador, Cordillera del Peuco, 1886, R.A.Philippi (SGO). 

The obvious type of the name L. chilensis was destroyed in B but details were depicted and 

described by F.G.P.Buchenau, Junc. S.-Amer. 418 (1879). The seed size, 6 stamens, and 

inflorescence shape clearly identify L. chilensis. The type of L. rigida does not bear 

Philippi's annotation of the name but the detailed description (with a number of measurements 

and individual features) allows identification of the type and its equation with 

L. chilensis. 

39. Luzula leiboldii Buchenau, Junc. S.-Amer. 418 (1880) 

Juncoides leiboldii (Buchenau) Kuntze, Revis. Gen. Pl. 2: 734 (1891). T: Chile, F.E.Leibold 2969; holo: W, 

not located; iso: S. 


Perennials 12–30 cm tall, densely caespitose, hairy; base thick, with extravaginal erect 

shoots; stolons absent. Basal leaves usually 5–10 cm long, 3.5–5.0 mm wide; cauline leaves 

1 or 2, 3.5–6.0 cm long, 2.0–3.0 mm wide; all leaves acute, rarely subobtuse, with margins 

±smooth to sparsely papillose, sparsely ciliate. Lower and middle bracts castaneousmembranous, 

with prominent veins, aristate, ciliate, up to 1.5 cm long; upper bracts c. 5 mm, 

similar to tepals. Inflorescence erect to suberect, rarely slightly nodding, usually interrupted 

at base, ‘echinate’ with erecto-patent acute bracts, usually 1.5–2.0 (–3.0) cm long, usually of 

3–7 few-flowered clusters subtended by conspicuous middle bracts. Bracteoles acuminate, 

membranous, 2.5–3.0 mm long. Tepals glabrous to sparsely ciliate, pale to castaneous brown 

at base, membranous-scarious above, acuminate-aristate, ±unequal; outer tepals 3.5–4.8 mm 

long; inner tepals 2.7–3.7 mm long. Stamens 6; anthers (0.7–) 0.8–1.2 (–1.5) mm long; 

filaments 0.4–0.6 mm long; style 0.5–0.6 mm long; stigmas 1.5–2.0 mm long. Capsule ovoid, 

subacute, deep castaneous, shorter than perianth; capsule segments 2.4–2.9 mm long. Seeds 

[unripe] 1.5–1.7 mm long, c. 1 mm wide; appendage indistinct, c. 0.1 mm long, fibrillate. 

Recorded from Santiago province, Chile, and Mendoza and Neuquén, Argentina. 85: AGS, 

AGW, CLN. Habitat unknown. Map 69. 

85. ARGENTINA NORTHWEST: Mendoza, Calmucó, A.Burkart et al. 13849 (SI13849). ARGENTINA 

SOUTH: Neuquén, Lacar, Cordón del Cerro Colorado (Between Lago Lacar and Lago Lolog), 25 Jan 1986, 

L.Bernardello & Moscone 620 (CORD). CHILE NORTH: Santiago, Rio Yeso, Lago Negro, 2700 m, 

W.Biese 937 (LIL); ‘Cordillera de Santiago’, R.A.Philippi (SGO, BM); Santiago, Cerro de Ramón, 2650 m, 

C.Ysandino (SGO). 

Closely related to L. chilensis, and perhaps only an infraspecific taxon of it. Luzula leiboldii 

is distinct in having longer tepals, longer anthers with higher anther/filament ratio, and tepals 

sometimes hairy. The material available is too scanty for evaluation of variation ranges. 

40. Luzula racemosa Desv., J. Bot. (Desvaux) 1: 162, tab. 6, fig. 3 (1808) 

Juncoides racemosa (Desv.) Kuntze, Revis. Gen. Pl. 2: 734 (1891). T: ‘in America calidiore’; lecto: P, 

designated here [left hand specimen, the rest is a fragment of a Luzula of another section]. 

Luzula spicata var. interrupta E.Mey., Linnaea 22: 415. 1849, non Luzula interrupta Desv. T: Mexico, 

Palyas de Jorullo, A.Humboldt; syn: n.v.; Mexico, Toluca, A.Humboldt; syn: n.v.; Mexico, Mt Orizaba, 

H.G.Galeotti; syn: n.v.; C.Schiede; syn: n.v. 

Luzula humilis Buchenau, Abh. Naturwiss. Vereine Bremen 4: 125 (1874); Luzula racemosa f. humilis 

(Buchenau) Buchenau, Bot. Jahrb. Syst. 12: 133 (1890); Luzula racemosa var. humilis (Buchenau) T.C.E.Fr., Nova 

Acta Regiae Soc. Sci. Upsal., ser. 4, 1: 167 (1905). T: Bolivia, La Paz, Larecaja, near Sorata, 3700–4200 m, 

G.Mandon 1448; lecto: GOET, fide H.Balslev, Fl. Neotrop. Monogr. 68: 67 (1996); isolecto: GH, P, S. 

77

78 


Luzula macusaniensis Steud. ex Buchenau, Abh. Naturwiss. Vereine Bremen 4: 131 (1874); Juncoides 

macusaniensis (Buchenau) Kuntze, Revis. Gen. Pl. 2: 734 (1891).. T: Peru, Puno, Macusani, 1854, 

W.Lechler 1839; lecto: P, fide H.Balslev, Fl. Neotrop. Monogr. 68: 467 (1996) (as ‘holotype’); isolecto: BR, 

G, GOET, K, P, S, WRSL. 

Illustrations: N.A.Desvaux, J. Bot. (Desvaux) 1: 162, tab. 6, fig. 3 (1808); H.Balslev, Fl. Neotrop. Monogr. 

68: 66, fig. 34, C, D (1996). 

Perennials (5–) 10–35 (–40) cm tall, caespitose, hairy; rhizome short, erect or ascending; 

stolons absent. Stem thin, nodding in upper part. Basal leaves canaliculate or flat, 5–18 cm 

long, typically 2.0–2.5 (–4.0) mm wide; cauline leaves (0–) 1 or 2, usually remote, narrow, 

usually 3.5–7 cm long; all leaves acute, with margins papillose-serrulate. Lower bract 

usually herbaceous, very narrow, up to 4–5 cm long, shorter than inflorescence; bracts 

supporting flower clusters linear-lanceolate, pale brownish-membranous, ciliate, up to 1 cm long, 

usually shorter than clusters. Inflorescence nodding, spike-like, (1.0–) 1.5–5 × 1.0–1.5 cm, 

of 3–8 elongated clusters, ±interrupted (rhachis visible), rarely compactly congested in high 

mountain forms (then only 1–1.5 cm long). Bracteoles ±membranous, 1.2–1.5 (–2.0) mm 

long, ovate, acuminate, ciliate. Tepals ±unequal, ±glabrous (rarely with few teeth or hairs), 

dark castaneous brown with paler apex and margins; outer tepals usually 2.0–2.4 (–3.0) mm 

long, lanceolate, acuminate-aristate; inner tepals 1.7–2.1 (–2.4) mm long. Stamens 3, rarely 

4–5; anthers 0.3–0.5 mm long; filaments 0.3–0.6 mm long; style 0.1–0.2 mm long; stigmas 

0.6–0.9 mm long. Capsules oblong-ovoid, usually ±obtuse, shorter than perianth, paler 

brown; capsule segments usually 1.5–1.7 (–1.8) × 0.9–1.1 mm. Seeds pale brown, shining, 

with darker apex, (0.7–) 0.9–1.0 mm long, c. 0.6 mm wide; appendage distinct, yellowish, 

0.1–0.2 mm long, sparsely short fibrillate. 2n=24, fide J.H.Beaman, Amer. J. Bot. 49(1): 

41–50 (1962), n.v. Fig. 18. 

Central (C Mexico and Guatemala) and South America, from Venezuela to Argentina and 

Chile. 79: MXC, MXG, MXS. 80: GUA, SMX. 82: VEN. 83: BOL, CLM, ECU, PER. 

85: AGS, AGW, CLN, CLS. Sunny and rocky sites in open woodlands and páramo, usually 

between 3000 and 4000 m. Map 70. 

79. MEXICO: Puebla, N side of Popocatépetl, J.H.Beaman 1742 (DUKE, GH, TEX, US). 80. GUATEMALA: 

San Marcos, Totonicapan, Tecum Uman ridge, 3340 m, J.H.Beaman 4180 (DUKE, NY, UC). 82. VENEZUELA: 

Mérida, Libertador, Sierra Nevada, Loma Redonda, 4045 m, L.J.Dorr & L.C.Barnett 5213 (AAU, NY). 

83. BOLIVIA: La Paz, Inquisivi, source of Rio Ocsalla, 3800 m, M.Lewis 40912 (AAU, LPB, MO). 

COLOMBIA: Páramo Choachi, 3000 m, A.Lindig 1404 (P, S). ECUADOR: Cotopaxi, 4 km S of Cotopaxi 

railway station, 3500 m, H.Balslev 1183 (AAU, NY). PERU: Huaráz, Huascarán, Quebrada Churup, 

3890–4260 m, D.N.Smith, Valencia & A.Gonzales 9629 (AAU, MO). 85. ARGENTINA NORTHWEST: 

Jujuy, Santa Ana, A.Burkart & N.S.Troncoso (SI11763); Tucumán, Ciénaga Grande, Cumbres Calchaquies, 

3800 m, S.Halloy s.n. (LIL567036). ARGENTINA SOUTH: Neuquén, Los Lagos, San Martín de los Andes, 

A.E.Rasp 6 (SI) 

An extremely variable species in most features of general habit. In various regions plants 

have features blurring the otherwise distinct limits of the species. In Mexico, flowers are 

close to the upper size limit, in Ecuador, an intermediate between L. racemosa and 

L. chilensis was found (Sayausí, G.Harling 1424, S). Plants in Argentina have a tendency 

towards longer peduncles of lower clusters, a feature found in related species. The plasticity 

of vegetative traits, depending often on available moisture and altitude, is also enormous. 

The whole complex is in a need of a detailed cultivation and karyological study. 

41. Luzula peruviana Desv., J. Bot. (Desvaux) 1: 160 (1808) 

Juncoides peruviana (Desv.) Kuntze, Revis. Gen. Pl. 2: 725 (1891). T: ‘in Terra Patagonala et montibus 

frigidetis Peruviae Pumbo, Antisana et Soluca.’; America calidiore, sine coll.; lecto: P, fide H.Balslev, Fl. 

Neotrop. Monogr. 68: 67 (1996) 1996: 67 [as holotype, left hand specimen typical]; Ecuador, Antisana, 

A.Humboldt; para: B-WILLD. 

Luzula boliviensis Buchenau, Abh. Naturwiss. Vereine Bremen 4: 128 (1874); Juncoides boliviensis 

(Buchenau) Kuntze, Revis. Gen. Pl. 2: 724 (1891). T: Bolivia, La Paz, Larecaja, Sorata, between Pongo and 

Anilaya, 3800 m, G.Mandon 1454; lecto: GOET, fide H.Balslev, Fl. Neotrop. Monogr. 68: 67 (1996); isolecto: 

BR, G, NY, P, S, UPS, US [p.p.; a few plants belong to L. racemosa or to intermediates]. 

Luzula peruviana var. longispica Barros, Darwiniana 10: 346 (1953). T: Argentina, Jujuy, Volcán, Loma 

del Tambo, 2500–3000 m, 22 Feb 1924, Schreiter 2909; holo: LIL.


Illustration: H.Balslev in Fl. Ecuador 11: 17, fig. 5 (1979). 

Perennials up to 60 cm tall, caespitose, hairy; rhizome vertical or ascending; stolons absent. 

Stem thick, rigidly erect, with only the uppermost part nodding. Basal leaves numerous, 

often squarrose, rigid, flat or canaliculate, usually 5–10 cm long and 2–4 mm wide; cauline 

leaves 2–4, densely hairy, to 13 cm long, evenly distributed on stem, often overlapping; all 

leaves acute, indistinctly papillose-serrulate. Lower bract herbaceous, densely hairy, up to 5 cm 

long, overtopping inflorescence; middle bracts supporting flower clusters purplish-brownish 

membranous, lanceolate, aristate, densely ciliate, c. 0.8–1.5 (–2.0) cm long, often exceeding 

the partial inflorescence. Inflorescence erect to slightly nodding, compact (rhachis usually not 

visible), usually 2.5–5 × c. 1.5 cm, often lobulate, rarely interrupted at base, dark castaneous 

brown, composed of c. 3–8 dense clusters. Bracteoles castaneous-membranous, ovatelanceolate, 

aristate, 2.0–2.5 mm, long ciliate. Tepals subequal or irregularly unequal, lanceolate, 

acuminate and aristate, usually ciliate, dark castaneous brown; upper part membranous with 

prominent midrib; longest tepals in flower 3.0–4.4 mm long. Stamens 3; anthers 0.3–0.4 mm 

long; filaments 0.6–0.9 mm long; style 0.1–0.2 mm long; stigma 0.5–1.1 mm long. Capsule 

oblong-ovoid, subacute to obtuse, paler brown; capsule segments 1.8–2.2 × 1.3–1.4 mm. 

Seeds paler brown, with darker apex, ellipsoidal, 1.1–1.4 mm long, c. 0.7 mm wide; 

appendage scarcely visible, c. 0.1 mm long, densely long fibrillate (seeds difficult to release 

from capsule). 

Centred to Ecuador and Peru, rare in Colombia, Bolivia and northernmost Argentina, with an 

isolated station in C Mexico; further exploration needed. 79: MXC. 83: BOL, CLM, ECU, 

PER. 85: AGW. In páramos, grasslands, rocky slopes, usually between 3800 and 4500 m. 

Map 71. 

79. MEXICO CENTRAL: Tlaxcala, Malinche, 4400–4450 m, J.H.Beaman 2223 (UPS). 83. BOLIVIA: La 

Paz, Murillo, Laguna Zongo, 4740 m, J.Brandbyge 582 (AAU). ECUADOR: Chimborazo, Mt Chimborazo, 

4550 m, E.Asplund 7938 (S, UPS); Pichincha, Páramo de Guamani, 3800–4100 m, H.Balslev 1618 (AAU); 

Imbabura, Volcán Cotocachi, 3900–4100 m, S.Laegaard 54498 (AAU). PERU: Puno, Araranca, 

4100–4300 m, F.W.Pennell 13456 (S); Huaron, 4500 m, J.F.Macbride & F.Featherstone 1132 (S); Ancash, 

Carhuaz, Quebrada Ulta, 4400–4600 m, D.N.Smith 11338 (AAU, MO). 85. ARGENTINA NORTHWEST: 

Salta, Santa Victoria, Cochita, Ruta 5, foot of cerro Campanario, 30-35 km W of Santa Victoria, 4100–4200 m, 

L.J.Novara 6683 (MCNS); Tucumán, Cumbres Calchaquíes, Callejones, 18 Feb 1915, L.Castillón (LIL 50636). 

Variable, and in many features similar to L. racemosa. Main diagnostic characters are its 

ciliate longer tepals, longer seeds with smaller, densely fibrillate appendage, and the habit of 

the inflorescence. Flowers of both species indicate autogamy but intermediates of unknown 

nature are known from Ecuador and Peru. 

42. Luzula vulcanica Liebm., Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 2: 

44 (1850) 

T: Mexico, Veracruz, Mt. Orizaba, 4250 m, F.Liebmann; lecto: C, fide H.Balslev, Fl. Neotrop. Monogr. 68: 65 

(1996) [as ‘holo’]; isolecto: C, GH, HAL, K, P, S. 

Illustration: H.Balslev, Fl. Neotrop. Monogr. 68: 66, fig. 34, A, B (1996). 


Perennials c. 10–45 cm tall, sparsely hairy; rhizome ascending to erect; stolons usually long, 

ascending. Stems erect, often slightly curved near the top (in the opposite direction to the 

nodding inflorescence). Basal leaves 3–15 cm long, usually 2–4 mm wide; cauline leaves 

usually 2, usually in upper half of stem, 1–5 (–11) cm long; all leaves acute, with papilloseserrulate 

margins. Bracts all scarious; lower bract membranous or castaneous (purplish), 

broadly lanceolate, c. 1 cm long, fimbriate-ciliate. Inflorescence a terminal spike like head, 

dark (reddish) castaneous brown, 1–4 × 0.5–1.2 cm, usually of 2–5 clusters, congested to 

slightly interrupted, suberect, if nodding then only the inflorescence not the upper part of 

stem. Bracteoles 1.5–2.0 mm long, usually castaneous-membranous. Tepals dark, reddishblackish, 

subequal, glabrous, usually 2.7–3.7 mm long, acuminate. Stamens 3; anthers 

0.4–0.5 mm long; filaments 0.8–1.0 mm long; style 0.1–0.2 mm long; stigmas c. 0.5–0.8 mm 

long. Capsule oblong-ovoid, ±trigonous above, paler castaneous; capsule segments 1.6–1.9 × 

79

80 


1.0–1.2 mm, finely rugose. Seeds ellipsoidal, paler brown, 0.9–1.0 mm long, 0.5–0.6 mm 

wide; appendage c. 0.1 mm long, fibrillate. Fig. 18. 

C Mexico and from Colombia to Bolivia. 79: MXC, MXG. 83: BOL, CLM, ECU, PER. 

Grasslands and dry open slopes in high altitudes. Map 72. 

79. MEXICO CENTRAL: Mexico, SW of Mt Ixtaccihuatl, 3850 m, J.H.Beaman 3527 (US). MEXICO 

GULF: Veracruz, Cofre de Perote, 4000 m, J.Dorantes 338 (F). 83. BOLIVIA: La Paz, Murillo, La Cumbre, 

4600 m, E.Asplund 4380 (UPS); Huayna Potosí, Lago Zongo, 4700 m, H.Balslev & S.G.Beck 1148A (AAU, 

LPB, NY). ECUADOR: Imbambura, Volcán Cayambe, 4100 m, P.C.D.Cazalet & T.D.Pennington 5723 (K, 

NY, US); Cotopaxi, Laguna Salayambo Cocha, 3800–4050 m, B.Øllgaard & H.Balslev 9891 (AAU); Azuay, 

road between Soldados and Angas, 3850 m, H.Balslev 1525 (AAU). PERU: Puno, Huancane, Occo Pampa, 

3125 m, R.S.Shepard 62 (GH, NY). 

43. Luzula antarctica Hook.f., Fl. Antarct. 1: 359 (1847) 

Juncoides antarctica (Hook.f.) Macloskie, Rep. Princeton Univ. Exp. Patagonia, Bot. 8: 302 (1904). T: Chile, 

Cape Horn, Hermite Is., J.D.Hooker 14; holo: K. 

Perennials 7–13 cm tall, densely caespitose, densely to sparsely hairy; rhizome short, 

vertical; stolons absent, plant base greyish straw-brown. Upper part of stem 0.3–0.4 mm in 

diam., arcuate. Basal leaves usually 3–5 cm long, ±flat to canaliculate, c. 2.0–3.0 (–3.5) mm 

wide, acute; margins subglabrous to densely ciliate, smooth or very remotely papillose. 

Cauline leaves (1–) 2, 1.5–3.5 cm long, c. 1.5 mm wide, acute. Lower bract (or uppermost 

leaf) usually slightly remote from inflorescence, brownish-membranous (or ±herbaceous), 

hairy, narrow, less than 2 cm long; upper bracts narrowly triangular to lanceolate or ovatelanceolate, 

dark castaneous brown, paler above, c. 0.7–1.0 cm long. Inflorescence small, 

1.0–1.5 × 0.7–0.9 cm, nodding, compact, dark castaneous brown, few-flowered. Bracteoles 

±lanceolate to narrowly triangular, castaneous brown to membranous-brownish, paler near 

apex, usually 2.5–4.0 mm long. Tepals unequal; inner tepals 2.8–3.4 mm long; outer tepals 

3.8–4.7 mm long, lacerate ciliate, dark castaneous brown (including bases of ciliae), 

lanceolate, acuminate, aristate (with a filiform tip). Stamens (0–2) 3; anthers 0.3–0.6 mm 

long; filaments 0.7–0.8 mm long; style c. 0.2 mm long; stigmas 1.0–1.2 mm long; gynoecium 

in some flowers rudimentary, but in most there are rudiments of 1 (–2) stamens. Capsule 

dark brown, ovoid to subglobular, subobtuse, shorter than perianth; capsule segments usually 

2.0–2.3 × 1.1–1.2 mm. Seeds ellipsoidal, brown; seed-coat not distinct, 1.0–1.1 mm long, 

0.6–0.7 mm wide; appendage yellowish, 0.10–0.15 mm long, fibrillate. 

Tierra del Fuego and adjacent territories, rare. 85: AGS, CLS. Exposed rocky slopes, 

grasslands. Map 73. 

85. SOUTH ARGENTINA: Tierra del Fuego, Cerro Almanza, Lashifashaj, 900–1000 m, D.M.Moore 1415 (H, K); 

Ushuaia, H.Roivainen 2321 (H). CHILE SOUTH: Tierra del Fuego, Río Azopardo, 600 m, 9 Mar 1896, 

P.Dusén 652 (UPS); 3 Mar 1908, C.J.F.Skottsberg 849 (S); Isla Desolación, Puerto Angusto, 300–400 m, 

8 Apr 1896, P.Dusén 709 (UPS); Sierra Valdiviesa, Río Betbedero, 11 Mar 1908, C.J.F.Skottsberg s.n. 

(SGO, UPS). 

Usually compared or synonymized with L. alopecurus on the basis of shared irrelevant 

characters [see D.M.Moore & M.C.Doggett, Bol. Soc. Argent. Bot. 17: 150–154 (1976)], but 

closer to the L. racemosa group. Distinct from the former in having three stamens and dark, 

nodding inflorescence, from the latter it can be distingushed by the compact inflorescence 

and much longer tepals. 

44. Luzula ruiz-lealii Barros, Darwiniana 10: 346 (1953) 

T: Argentina, Mendoza, Tunuyán, Valle del Alto Tunuyán, 16 Feb 1934, A.Ruíz Leal 2140; holo: LIL46295. 

Luzula parvula Barros, Darwiniana 10: 356 (1953). T: Argentina, Mendoza, San Carlos, Laguna del 

Diamante, 3300 m, 16 Jan 1949, A.Ruíz Leal 11717; holo: LIL. 

Illustrations: M.Barros, Darwiniana 10: 343, fig. 19 & 357, fig. 25 (1953). 

Perennials to 15 cm tall, dwarf, densely caespitose, cushion-forming, subglabrous to sparsely 

hairy; rhizome ±erect, ±many-headed; stolons absent, with base covered with numerous 

brownish remains of leaf bases. Basal leaves rigid, glabrescent or ciliate, thick, canaliculate,


Figure 18. Luzula vulcanica. A, habit, note stolons; B, flower. L. racemosa. C, habit, note 

lack of stolons; D, flower. (A, B, B.Øllgaard & H.Balslev 9891, AAU; C, D, P.R.Bell 78, A). 

Scale bars: A, C = 5 cm; B, D = 2 mm. Drawn by K.Tind Originally published in G.Harling 

& B.Sparre (eds.), Fl. Ecuador 11: 15, fig. 4; 17, fig. 5 (1979), reproduced with permission, 

from H.Balslev, Fl. Neotrop. Monogr. 68: 66, fig. 34 (1996), © Lunds Botaniska Förening, 

© 1996, The New York Botanical Garden. 

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82 


3–4 cm long, c. 1 mm wide, with ±obtuse to subobtuse tip and margins ±smooth; cauline 

leaves absent. Basal bract pale straw-brownish membranous, oblong-lanceolate, acute, 

ciliate, up to 1 cm long. Inflorescence spike-like, densely congested, terminal, cylindricalovoid,1.0–1.5 

× c. 0.7–0.8 cm, of 2–6 few-flowered clusters, or rarely of 1–3 remote basal 

clusters. Bracteoles whitish scarious, ovate, acuminate, ciliate, to 1.5–1.8 mm long. Tepals 

glabrous, ±equal, pale straw-brown or membranous above, brown below, acuminate 

(subaristate), with inner tepal ±mucronate, usually 2.1–2.6 mm long. Stamens 3–5 (–6); 

anthers 0.4–0.6 mm long; filaments 0.4–0.6 mm long; style c. 0.1 mm long; stigmas to 1 mm 

long. Capsule ovoid, subacute, paler brown, shorter than perianth; capsule segments 1.6–1.8 × 

c. 1.0 mm. Seeds 1.0–1.1 mm long, c. 0.6 mm wide; appendage yellowish, c. 0.2 mm long, 

fibrillate. 

Endemic to Mendoza, Argentina. 85: AGW-ME. Open patches in dry grasslands, above 

3000 m. Map 74. 

85. ARGENTINA NORTHWEST: Mendoza, San Carlos, Laguna del Diamante, 3200 m, 16 Jan 1952, A.Ruíz 

Leal 14584 (SI); 16 Jan 1949, A.Ruíz Leal 11717 (LIL264113, MERL); Mendoza, Tunuyán, Valle del Alto 

Tunuyán, Cerro Pan de Azúcar, 6 Feb 1934, A.Ruíz Leal 2140 (LIL, MERL). 

A peculiar dwarf form, superficially similar to L. ulophylla of New Zealand. Number of 

stamens varies within inflorescence and cannot be used to separate L. ruiz-lealii and 

L. parvula. Suspected to be a depauperate form parallel to that of L. racemosa 

(L. macusaniensis) but cannot be equated with any of the southern hexandrous forms. 

Doubtful names: 

Luzula psilophylla Phil., Linnaea 33: 267 (1865) 

Luzula chilensis f. psilophylla (Phil.) Fuentes, Bol. Mus. Nac. Hist. Nat. 90: 138 (1917). T: In Andibus 

‘Talcaregue’. [J.Gay]; holo: n.v. 

The type material has not been located. The protologue description does not give relevant 

information, and later herbarium material annotated by Philippi is taxonomically 

heterogenous, including Luzula leiboldii, L. chilensis and L. brachyphylla. 

Luzula chilensis var. pallida Buchenau ex Fuentes, Bol. Mus. Nac. Hist. Nat. 90: 138 (1917) 

T: [not indicated]. 

Presumed Hybrids 

Luzula ×somedana Fern.-Carv. & Fern.Prieto in J.A.Fernández Prieto & M. de C.Fernández- 

Carvajal, Stud. Bot. 2: 134 (1983) = Luzula pediformis (Chaix) DC. × L. sylvatica subsp. 

henriquesii (Degen) Pirajá 

T: Spain, Asturias, Somiedo, Sierra del Parámo, Laguna Cavera, 1720 m, 21 Jul 1978, J.A.Fernández-Prieto; 

holo: FCO8195; see also J.A.Fernández Prieto & C.Fernández-Carvajal, Stud. Bot. 2: 136, fig. A, B, C (1983). 

A seed-sterile hybrid. 

Luzula ×gayana Font Quer & Rothm., Cavanillesia 7: 175 (1936) = Luzula pediformis 

(Chaix) DC. × L. caespitosa (E.Mey.) Steud. 

T: Spain, Asturias, Cueto de Arvas, 1700 m, 15 Jul 1935, P.Font Quer & W.Rothmaler; holo, n.v. 

Material not seen but the existence of the hybrid is probable. 

Luzula ×marceloi Rivas Mart., Anales Inst. Bot. Cavanilles 21(1): 289 (1963), nom. inval. = 

Luzula caespitosa (E.Mey.) Steud. × L. spicata (L.) DC. 

T: Spain, Sierras de Guadarrama y Gredos, Ladera del Noruego, 2200 m, S.Rivas Martínez; holo: n.v.


Sect. 6. THYRSANOCHLAMYDEAE 

Luzula sect. Thyrsanochlamydeae Satake, in T.Nakai & M.Honda, Nov. Fl. Jap. 1: 30 (1938) 

T: Luzula unalaschkensis (Buchenau) Satake; lecto, designated here. 

Luzula subsect. Thyrsanochlamydeae Novikov, Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol., 95(6): 67 

(1990), nom. illeg. T: Luzula confusa Lindeb. 

Luzula subsect. Tundricolae Kovt., Bot. Žurn. (Moscow & Leningrad) 72: 1399 (1987). T: Luzula 

tundricola Gorodkov ex V.N.Vassil. 

Luzula subsect. Confusae Kovt., Bot. Žurn. (Moscow & Leningrad) 72: 1399 (1987). T: Luzula confusa 

Lindeb. 

Luzula subsect. Nivales [Chrtek & Křísa ex] Kovt., Bot. Žurn. (Moscow & Leningrad) 72: 1399 (1987). 

T: Luzula nivalis (Laest.) Spreng. 

Perennials. Leaves with tip acute, acuminate or mucronate. Lower bract short, membranous 

to brownish-membranous. Inflorescence subcongested, or in glomerules on thin peduncles 

which are usually arcuate or nodding, less often straight. Seeds ellipsoidal to narrowly 

ellipsoidal; appendage basal, short, up to 0.2 mm long, or lacking; base fibrillate. 

Five species, almost confined to the Arctic and rare in the mountains in northern temperate 

regions. They represent a link between sect. Diprophyllatae and the type section. 

In particular, L. arcuata and L. confusa may have an alloploid origin of the above parentage, 

as indicated by their karyotypes (Nordenskiöld, 1949, 1951). 

1 Basal leaves usually 4.5–6.0 mm wide, bluish green; plants usually tall, 

20–40 cm tall [California] 45. L. subcongesta 

1: Basal leaves usually narrower than 4 mm; plants usually less than 20 cm 

tall [circumpolar but north of California] 

2 Basal leaves with obtuse, often slightly swollen tip with a spinuliform 

mucro; seed base with appendage c. 0.2 mm long and fibrillate 49. L. nivalis 

2: Basal leaves subabruptly to gradually narrowing to acute or acuminate 

tip; seed base with short (to 0.1 mm long) appendage or without 

appendage, and always fibrillate 

3 Plant base straw-brown to dull brownish; flower clusters pale brown 

to straw-brown, ellipsoidal 46. L. kjellmaniana 

3: Plant base brown to purplish or reddish, usually shining; flower 

glomerules or clusters dark to blackish brown, ±globular or ellipsoidal 

4 Peduncles c. 0.2 mm in diam., with secondary branches, usually 

arcuate to nodding; glomerules (1–) 2–6 (–8)-flowered, broader than 

wide; number of glomerules usually 5–10; stem less than 0.5 mm in 

diam. below inflorescence 47. L. arcuata 

4: Peduncles c. 0.3 mm in diam., without secondary branches, usually 

straight, erect, less often arcuate; subsessile clusters more than 

10-flowered; number of clusters usually 1–3; stem 0.5–0.9 mm in 

diam. below inflorescence 48. L. confusa 

45. Luzula subcongesta (S.Watson) Jeps., Fl. Calif. 1: 258 (1921) 

Luzula spadicea var. subcongesta S.Watson, Bot. California 2: 202 (1880); Luzula parviflora var. 

subcongesta (S.Watson) Buchenau, Bot. Jahrb. Syst. 12: 110 (1890); Juncoides parviflora var. subcongesta 

(S.Watson) E.Sheld., Bull. Geol. Nat. Hist. Surv. 9: 63 (1894); Juncoides subcongesta Coville, Muhlenbergia 

1: 105 (1904). T: California, Sierra Nevada, near Donner Lake, J.Torrey; syn: n.v. 

Illustration: Jepson, Fl. Calif. 1: 259, fig. 45c, d, e (1921). 

Map: J.Coffey Swab, Fl. North Amer. 22: 267 (2000). 

Perennials, loosely tufted, subglabrous, bluish green, with horizontal rhizome. Stem 20–40 cm 

long; cauline leaves 3–4, usually 3–5 cm long, 3–6 mm wide; basal leaves up to 10 (–15) cm 

83

84 


long, usually 4.5–6 mm wide; all leaves with distinctly acute tip, papillose-serrulate margins 

and sparse hairs near sheath. Inflorescence bracts straw-membranous or with reddish tinge; 

lower bracts up to 1.5 cm long, ciliate. Inflorescence up to 4–6 cm long, corymbose with 

4–20 glomerules on capillary arcuate peduncles c. 0.2 mm in diam., each distally branched; 

glomerules usually 3–7-flowered. Bracteoles usually c. 1.5 cm long, ciliolate. Tepals equal, 

1.7–2.1 mm long, 0.5–0.6 mm wide, ovate-lanceolate, acute, entire, dark brown. Stamens 6; 

anthers 0.3–0.6 mm long; filaments 0.5–0.7 mm long; style c. 0.3 mm long; stigmas c. 1.0 mm 

long. Capsule ovoid with subconical top, shorter than tepals; capsule segments dark brown, 

lustrous, 1.6–1.8 × 0.9–1.1 mm. Seeds narrowly ellipsoidal, 1.1–1.2 × 0.6 mm, medium 

brown, distally apiculate, with fibrillate base; appendage missing. 2n=24 [BL], 

H.Nordenskiöld, Hereditas 37: 330–331, 1951. Fig. 19. 

Endemic to California. 76: CAL. Gravelly stream and lake banks, grassy patches among 

rocks, usually at or above 3000 m. Map 75. 

76: CALIFORNIA: Sierra Nevada, Tuolumne Co., head of Virginia Canyon, c. 3200 m, C.W.Sharsmith 4148 

(DS, K, S, UC); Tulare Co., Rock Ck, c. 3400 m, J.T.Howell 25567 (CAS, S); J.T.Howell 26216A (UPS); 

Mono Co., Hall Natural Area, c. 3000 m, A.R.Moldenke & H.N.Moldenke 24653 (AAU); Mono Co., Cascade 

Lake, NE of North Peak, 3300 m, J.Clausen 1934 & A.R.Kruckeberg (K); Shasta Co., Mt. Shasta, W.H.Brewer 

1378 (GH, UC, US); Placer Co., Donner Pass, c. 2400 m, A.A.Heller 7135 (K). 

Closely similar to species of the sect. Diprophyllatae, both in seed and leaf characters, but 

the inflorescence is composed of dense glomerules. 

46. Luzula kjellmaniana Miyabe & Kudo, Trans. Sapporo Nat. Hist. Soc. 5: 38 (1913) 

Luzula arcuata f. latifolia Kjellm., in A.E.Nordenskiöld, Vega-Exped. Vetenskapl. Iagkttagelser 1: 566 

(1882), nom. illeg., non Laest.; Luzula confusa f. latifolia Buchenau, Bot. Jahrb. Syst. 12: 125 (1890); Luzula 

arctica var. latifolia (Buchenau) Nilsson ex Buchenau, in H.G.A.Engler, Pflanzenr. (iv.36) 25: 69 (1906); 

Luzula multiflora var. kjellmaniana (Miyabe & Kudo) Sam., in E.Hultén, Kongl. Svenska Vetenskapsakad. 

Handl., ser. III, 5(1): 227 (1927), nom. illeg.; Luzula beeringiana Gjaerev., Kongel. Norske-Vidensk. Selsk. 

Skr. (Trondheim) 1958 (5): 63 (1958), nom. illeg.; Luzula multiflora subsp. kjellmaniana (Miyabe & Kudo) 

Tolm., Arktic Fl. SSSR 4: 48 (1963); Luzula hyperborea var. latifolia (Buchenau) B.Boivin, Naturaliste 

Canad. 94: 526 (1967); Luzula arctica subsp. latifolia (Buchenau) Porsild, Nat. Canad., Bot. 4 [1974]: 15 

(1975), nom. illeg. T: Russia, Chukotka, St. Lawrence Bay, 20–21 Jul 1879, F.R.Kjellman; lecto: S, fide 

L.Hämet-Ahti, in L.Hämet-Ahti & Virrankoski, Ann. Bot. Fenn. 8: 157 (1971). 

Luzula tundricola Gorodkov ex V.N.Vassil., Bot. Mater. Gerb. Bot. Inst. Komarova Akad. Nauk SSSR 15: 40 

(1953). T: Russia, Anadyr, Novomariinsk, 1932, V. N. Vasilev 168; holo: LE. 

Illustrations: E.Hultén, Fl. Alaska 300 (1968); J.Coffey Swab, Fl. North Amer. 22: 264 (2000). 

Maps: E.Hultén & M.Fries, Atlas N. Eur. Vasc. Pl. 1: map 170 (1986), as L. tundricola; E.Hultén, Fl. Alaska 

300 (1968); N.K.Kovtonyuk, in L.I.Malyshev & G.A.Peshkova, Fl. Sibiri 4 (Araceae – Orchidaceae): map 43 

(1987) [Siberia]; J.Coffey Swab, Fl. North Amer. 22: 267 (2000). 

Perennials 8–20 cm tall; rhizome ascending, long, without stolons. Basal leaves 5–8 cm long, 

3–4 mm wide; sheath bases straw-brown to dull brownish. Cauline leaves 2(–3), usually 

2.5–4.0 cm long, up to 2.5 mm wide. All leaves ±flat with leaf tip subobtuse, often with a 

short acute mucro; margins papillose-serrulate. Lower bract short, membranous, brownish at 

base. Inflorescence composed of 1–2 sessile and 1–4 pedunculate clusters (these sometimes 

composed of a few smaller sessile subclusters, rarely whole inflorescence subcongested); 

clusters ellipsoidal, usually 6–7 × 4 mm, pale brown to straw-brown; peduncles capillary, 

erect to arcuate. Bracteoles usually 1.5–1.8 mm long, pale brownish to membranous, 

fimbriate-ciliate. Tepals subequal (outer longer by 0.2–0.4 mm), 2.0–2.6 mm long, medium 

brown with broad membranous to stramineous margin above, entire. Stamens 6; anthers 

0.4–0.5 mm long; filaments 0.5–0.7 mm long; style 0.2–0.3 mm long; stigmas c. 0.8 mm 

long. Capsule subacuminate, usually shorter than tepals; capsule segments ellipsoidal, 

brown, 1.5–1.6 × 1.0 mm. Seeds narrowly ellipsoidal, 0.9–1.0 × 0.5–0.6 mm, brown; 

appendage minute, less than 0.1 mm long; base fibrillate. 2n=36 [12'AL', 24'BL'], L.Hämet- 

Ahti & Virrankoski, Ann. Bot. Fenn. 8: 157, 1971. Fig. 20.


Figure 19. Luzula subcongesta. A, habit; B, flower cluster; C, flower; D, tepal; E, seed. 

L. confusa. F, habit; G, inflorescence; H, flower; I, tepal and stamen; J, seed. 

(A-E, J.T.Howell 25567, S; J.T.Howell 26216A, UPS; F-I, N.S.Vodop’yanova 301, PRA; 

W.E.Parry s.n., BM). Scale bars: A = 5 cm; B = 2 mm; C = 1 mm; D = 1mm; E, J = 0.5 mm; 

F = 5 cm; G = 1 cm; H, I = 1 mm. Drawn by © E.Smrčinová. 

85

86 


Northern regions of Siberia and the Russian Far East, Kamchatka, Alaska and NW Canada. 

30: KRA, YAK. 31: KAM, KHA, KUR, MAG. 70: ALU, ASK, NWT, YUK. 71: BRC. 

In drier tundra, and on rocky slopes. Map 76. 

30: KRASNOYARSK: Taymyr, Syndasko R., N.S.Vodop’yanova 412 (PRA). YAKUTIYA [SAKHA]: 

Bulkur, 1898, H.Nilsson s.n. (LD); Balganach, 1898, H.Nilsson s.n. (LD). 31: MAGADAN: Chukotka, 

15 km SSW of Provideniya, 23 Jun 1992, A.Svensson (S). 70: ALEUTIAN IS.: Akutan, Norberg 71 (S); 

Amlia Is., W.J.Eyerdam 1234 (BM). ALASKA: Alaska Range Distr., Dry Ck, c. 1100 m, A.Viereck & Jones 

5772 (S); Pribilof Is., St. Paul, E.Hultén 7459 (S); Brooks Range, De Long Mtns, Liberator Lake, K.Holmen & 

O.Mårtensson 61-1640 (UPS); Port Clarence, Teller Reindeer Station, B.A.Walpole 1963 (K). 

NORTHWEST TERRITORIES [Nunavut]: Mackenzie R. Delta, Kittigaznit Is., A.E.Porsild & R.T.Porsild 

2333 (S); Tungsten, Marris 11225 (BM). 

The accepted name has often been misinterpreted for plants of the sect. Luzula. For 

discussion, see L.Hämet-Ahti & Virrankoski, Ann. Bot. Fenn. 8: 157–158 (1971). 

47. Luzula arcuata (Wahlenb.) Sw., Summa Veg. Scand. 13 (1814) 

Juncus arcuatus Wahlenb., Fl. Lapp., p. 87, tab. 4, 1812, nom. conserv. propos. [nom. illeg., Juncus 

pediformis Chaix in syn.]; Juncoides arcuata (Wahlenb.) Druce, Bot. Soc. Exch. Club Brit. Isles 8: 85 (1929). 

T: Sweden, Piteå Lappmark på Örfjället, 6 Sep 1807, G.Wahlenberg; lecto: UPS, fide Moberg & Nilsson, 

Nord. J. Bot. 11: 293 (1991). [further authentic material: S, UPS, WU]. 


Perennials, sparsely hairy, with ascending rhizomes and short stolons. Leaves subabruptly 

narrowing to acute tip. Basal leaves usually up to 4 mm wide. Stem usually 0.3–0.5 mm in 

diam. Lower bract membranous-brownish, usually 3–8 (–15) mm long. Inflorescence 

corymbose, composed of 3–17 few-flowered glomerules usually as long as wide or wider; 

peduncles with secondary branches, capillary, usually c. 0.2 mm in diam., arcuate or 

nodding. Stamens 6. Seeds narrowly ellipsoidal; appendage absent or up to 0.15 mm long; 

base fibrillate. 

Arctic regions of Europe; the Arctic and mountains of N Asia; N America from Washington 

and British Columbia northwards; Japan, Korea and, probably, China. Rocky or gravelly 

places. Two subspecies are recognized. 

Basal leaves up to 2 mm wide; seed appendage lacking 

Basal leaves more than 3 mm wide; seed appendage developed, 

47a. subsp. arcuata 

to 0.15 mm long 47b. subsp. unalaschkensis 

47a. Luzula arcuata (Wahlenb.) Sw. subsp. arcuata 

Luzula arcuata f. capillaris Laest., Bot. Not. 1858: 144 (1858). T: Sweden, Pite Lappmark, "Njepusurte i 

Piteå Lappmark", L.L.Laestadius; syn: n.v.; [Norway, Finnmark] Koutokeino, L.L.Laestadius; syn: n.v. 

Illustration: J.Lid, Norsk & Svensk Fl. 203, fig. 100a (1963). 

Perennials up to 15 cm long, sparsely hairy to glabrescent, loosely caespitose, with branched 

creeping or ascending thin rhizome and ascending stolons. Basal leaves canaliculate, up to 

5–7 cm long, up to 1.5–2.0 mm wide, subabruptly narrowing to acute tip; plant base brownish 

to purplish, slightly lustrous; cauline leaves 1 (–2), usually 1.5–2.0 (–3.0) cm long, to 1.2 mm 

wide; stem thin, usually less than 0.5 mm in diam. below inflorescence. Lower bract 

membranous, often suffused brownish, 3–5 mm long, ciliate above. Inflorescence lax, 

corymbose, composed of (3–) 5–15 small glomerules; peduncles smooth, thin, c. 0.2 mm long, 

arcuate and nodding, up to 1.5–2 cm long, often branched (secondary branches usually 1–3); 

glomerules c. 4 × 3.5–6 mm, usually (1–) 2–5 (–8)-flowered, often with some flowers with 

visible peduncles. Bracteoles 1.0–1.2 mm long, ovate, brownish below, scarious and fimbriateciliolate 

above. Tepals ±equal; outer tepals acuminate; inner tepals ±mucronate or acute, deep 

brown, usually 2.0–2.2 mm long. Stamens 6; anthers 0.4–0.6 mm long; filaments c. 0.3 mm 

long; style 0.4–0.5 mm long; stigma 1.1–1.3 mm long. Capsule ovoid to ellipsoidal, slightly


Figure 20. Luzula arcuata subsp. arcuata. A, habit; B, flower glomerule; C, flower; D, tepal 

and stamen; E, seeds. Luzula kjellmaniana. F, habit; G, inflorescence; H, flower; I, tepal and 

stamen; J, leaf apex; K, seed. (A–J, J.Montell s.n., PR; F–K, A.E.Porsild & R.T.Porsild 

2333, S; E.Hultén 7459, S). Scale bars: A = 5 cm; B = 4 mm; C, H = 1 mm; D, I = 1 mm; 

E, K = 0.5 mm; F = 5 cm; G = 1 cm; J = 4 mm. Drawn by © E.Smrčinová. 

87

88 


mucronate, shorter than or equalling tepals; capsule segments ovate, 1.7–1.9 × 1.1–1.2 mm, 

dark brown above. Seeds brown to pale brown, ellipsoidal, 1.0–1.1 × 0.6 mm, apically 

darker, apiculate, basally fibrillate. 2n=36, 42, 48, cf. Dawe & Murray, Canad. J. Bot. 59: 

1373–1374 (1981); H.Nordenskiöld, Bot. Not. 1949: 85–86 (1949). Fig. 20. 

Subarctic and mountain regions of northern Europe, with a NW limit in Iceland and reaching 

northern European Russia and Novaya Zemlya in the NE. Reported to occur in Greenland but 

material not seen. 10: FIN, FOR, GRB, ICE, NOR, SVA, SWE. 14: RUN. 30: WSB. 

Mountain and subarctic tundra and stony slopes. Map 77. 

10: FØROYAR: Kunö Is., 24 Aug 1897, Hartz & C.E.Ostenfeld (LD, S). GREAT BRITAIN: Scotland, 

Easterness, Cairngorm, A.Melderis 425 (S). NORWAY: Dovre Fjeld, Fogstuen, 1000 m, 22 Jul 1889, 

K.G.Baenitz (W). SVALBARD: Safehaven, 30 Aug 1882, A.G.Nathorst s.n. (S); Jan Mayen, Wildberget, 

Lid [Fl. Exs. Ins. J.Mayen] 18 (LD). SWEDEN: Åsele Lappmark, Laxfjället, 12 Jul 1889, G.Halldin s.n. 

(LD); Lycksele Lappmark, Tärna socken, Brandsfjället, 24 Jul 1937, E.Asplund s.n. (LD); ‘ex alpibus 

Lapponiae Pitensis’ G.Wahlenberg s.n. (WU). 14. RUSSIA NORTH: Lapponia ponojensis, Ponoy, 27 Aug 

1899, J.Montell s.n. (H). 

Sometimes difficult to distinguish from L. confusa in Scandinavia because of frequent 

occurrence of intermediates of hybrid origin 

47b. Luzula arcuata subsp. unalaschkensis (Buchenau) Hultén, Ark. Bot. 7(1): 32 (1968) 

Luzula arcuata var. unalaschkensis Buchenau, Bot. Jahrb. Syst. 12: 124 (1890); Luzula unalaschkensis 

(Buchenau) Satake, in T.Nakai & M.Honda, Nov. Fl. Jap. 1: 31 (1938) & J. Jap. Bot. 14: 7 (1938). T: Alaska, 

Unalaschka, Mertens; lecto: W, designated here. [Unalaschka, J.F.G.Eschscholtz; orig. syn: LE, n.v.]. 

Luzula arcuata var. kamtschadalorum Sam., in E.Hultén, Kongl. Svenska Vetenskapsakad. Handl., ser. III, 

5(1): 223 (1927); Luzula kamtschadalorum (Sam.) Gorodkov ex Krylov, Fl. Zap. Sibiri 3: 551 (1929); Luzula 

unalaschkensis subsp. kamtschdalorum (Sam.) Tolm., Arktic Fl. SSSR 4: 40 (1963). T: Russia, Kamchatka, 

Avatcha volcano, 20 Aug 1920, E.Hultén 670; lecto: S, fide L.Hämet-Ahti, Ann. Bot. Fenn. 10: 123 (1973); 

isolecto: K. 

Luzula beringensis Tolm., Arktic Fl. SSSR 4: 39 (1963). T: Russia, Terra Koriakorum, prope sinum Korfii 

maris Beringii, 19 Aug 1960, A.I.Tolmachev, Stepanova & Fedorova; holo: LE. 

Illustrations: E.Hultén, Fl. Alaska 299 (1968); L.Hämet-Ahti, Ann. Bot. Fenn. 10: 126, fig. 3; 127, fig. 4 

(1973); V.S.Novikov, in S.S.Kharkevich, Sosud. Rast. Soviet. Dal. Vost. 1: 83, plate vi (1985). 

Maps: E.Hultén, Fl. Alaska 299 (1968); N.K.Kovtonyuk, in L.I.Malyshev & G.A.Peshkova, Fl. Sibiri 4 

(Araceae – Orchidaceae): map 34 (1987) [Siberia]; J.Coffey Swab, Fl. North Amer. 22: 266 (2000). 

Perennials 10–25 cm tall; rhizome ascending; stolons ascending, usually 2–3 cm long. Leaves 

±flat, sparsely ciliate, subabruptly narrowing to an acute tip; basal leaves usually 8–12 cm 

long, 3–4 (–5) mm wide; sheath base brown to purplish; cauline leaves 1–3, to 3–5 (rarely 8) cm 

long, 1.5–3.0 mm wide; sheaths sparsely ciliate above. Stem 0.3–0.4 (–0.5) mm diam. below 

inflorescence. Lower bract usually 5–8 (–15) mm long, brownish-membranous. Inflorescence 

usually corymbose with (2–) 6–10 (–17) glomerules; glomerules usually of (1–) 3–6 flowers 

each; peduncles capillary, arcuate, often nodding, often branched, usually reddish. Bracteoles 

±ovate-lanceolate, c. 1.5 mm long, brownish to membranous, ciliate. Tepals ±equal, usually 

2.0–3.0 × 0.6–0.8 mm, ovate to lanceolate, acuminate, often ciliolate, medium to pale brown. 

Stamens 6; anthers 0.4–0.6 mm long; filaments 0.4–0.5 mm long; style 0.3–0.4 (–0.5) mm 

long; stigmas 1.1–1.5 mm long. Capsule ellipsoidal with a short beak, shorter than tepals; 

capsule segments pale brown to brown, 1.9–2.2 × 1.0–1.4 mm. Seeds brown, narrowly 

ellipsoidal, 1.0–1.1 × c. 0.6 mm; appendage very short, up to 0.15 mm long; base fibrillate 

(on the top of caruncle). 2n=36, G.S.Knaben, Nytt Mag. Bot. 15: 244 (1968). 

Russian Far East, Japan, Baikal Lake region, northwestern America from Washington and 

British Columbia to Alaska and Nunavut [NW Territories]. Not confirmed but probably 

present in China and Mongolia. 30: BRY, CTA, IRK. 31: KAM, KUR, MAG. 38: JAP, KOR. 

70: ALU, ASK, NWT, YUK. 71: ABT, BRC. 73: WAS. Rocky or gravelly snow-patches, 

heaths and slopes, often on bare ground. Map 78. 

30: BURYATIYA: Baikal Range, sources of Kurkula R., T.V.Egorova & V.N.Siplivinsky s.n. (OULU). 

31: MAGADAN: upper Main R., 12 Aug 1932, B.N.Gorodkov & B.A.Tikhomirov s.n. (S). KAMCHATKA: 

Natchika, Mt. Tschopotschka [Shapochka], 4 Aug 1908, V.L.Komarov s.n. (S). KURIL IS.: Shumushu,


G.Koidzumi 39053 (TNS). 38: JAPAN: Nagano, Daitenjo-Lake, J.Sugimoto 13437 (KYO). KOREA: Mt. 

Pai-tou-shan, T.Nakai s.n. (TI). 70: ALEUTIAN IS.: Unalaska, E.Hultén 6671 (S). ALASKA: White 

Pass, 20 Aug 1929, S.Enander s.n. (UPS). YUKON: Canol Road, pass between Teslin R. and Nisutlin R., 

A.E.Porsild & A.Breitung 10983 (S). 71: BRITISH COLUMBIA: Red Rose Tungsten Mine, 15 km E of 

Skeena Crossing, c. 1600 m, J.A.Calder et al. 15173 (S). 

A variable subspecies; in Asia, towards the southeast, a gradual increase in perianth and 

capsule size can be observed, and plants from Japan often have solitary flowers on peduncles. 

The Japanese plants have often been referred to as L. wahlenbergii; literature records under 

the latter name from NE China may also refer to subsp. unalaschkensis. 

48. Luzula confusa Lindeb., Bot. Not. 1855: 9 (1855) 

Luzula arcuata f. confusa (Lindeb.) Kjellm., Vega-Exped. Vetenskapl. Iagttagelser 2: 59 (1883), n.v.; Luzula 

arcuata subsp. confusa (Lindeb.) O.C.Dahl, in A.G.Blytt, Handb. Norges Fl. 201 (1906); Juncoides confusa 

(Lindeb.) A.Heller, Cat. N. Amer. Pl., 3rd edn, 70 (1910). T: [Norway] Dovre, 1854, C.J.Lindeberg; lecto: 

S, designated here. 

Luzula spicata var. kjellmanii Nath., Kongl. Vetensk. Akad. Handl. 30: 28 (1884). T: N Greenland, Kap 

York, Insugigsok, 26 Jul 1883, A.G.Nathorst; syn: W. 

Luzula hyperborea R. Br., Chlor. Melvill. [Parry’s 1 st Voyage App.] 283 (1823), nom. rejic. prop.; Luzula 

arcuata var. hyperborea (R.Br.) Rink, in J.Reinhardt et al., Naturhist. Bidr. Grønl. 119 (1857); Juncoides 

hyperborea (R.Br.) Coville, Mem. Torrey Bot. Club 5: 107 (1894). T: Spitzbergen, W.Scoresby; syn: BM; 

Greenland, W.Scoresby; syn: BM; Melville Island, W.E.Parry; syn: BM, PH. [covering both Luzula confusa 

and Luzula nivalis]. 

Luzula confusa var. subspicata Lange, Consp. Fl. Groenland. 127 (1880); Luzula confusa f. subspicata 

(Lange) Krylov, Fl. Zap. Sibiri 3: 552 (1929). T: [Greenland] numerous syn: C. 

Luzula hyperborea var. extensa Scheutz, Kongl. Svenska Vetenskapsakad. Handl. 22: 169 (1888). T: [Russia, 

Taymyr] Nikandroviskij ostrov et Briochovskij ostrov; syn: n.v. 

Luzula confusa f. normalis Krylov, Fl. Zap. Sibiri 3: 552 (1929). T: [not indicated]. 

Illustrations: J.Lid, Norsk & Svensk Fl. 203, fig. 100b (1963); E.Hultén, Fl. Alaska 301 (1968). 

Maps: E.Hultén & M.Fries, Atlas N. Eur. Vasc. Pl. 1: map 169 (1986); E.Hultén, Fl. Alaska 301 (1968); 

N.K.Kovtonyuk, in L.I.Malyshev & G.A.Peshkova, Fl. Sibiri 4 (Araceae – Orchidaceae): map 35 (1987) 

[Siberia]. 

Perennials, usually 10–15 (–20) cm tall, sparsely hairy, loosely to densely caespitose with 

ascending densely branched to simple rhizome, sometimes with short ascending vegetative 

shoots; plant base usually deep purple-brown, shiny. Basal leaves up to 6–9 cm long, 

±subcanaliculate to ±flat, 1.5–2.5 mm wide; tip acuminate; cauline leaves 1–2, usually 2–4 cm 

long, 1.0–1.5 mm wide. Stem 0.5–0.9 mm in diam. below inflorescence. Lower bract 

membranous, less often brownish-membranous, usually 4–9 mm long. Inflorescence 

composed of one sessile and often 1-2 pedunculate clusters; peduncles without secondary 

branches, erect, straight, less often arcuate or recurved, c. 0.3 mm in diam.; sessile cluster 

bigger (sometimes with a discernible basal subcluster), usually more than 10-flowered, 7–10 × 

6–7 (–9) mm; pedunculate clusters smaller, 5–12 (–14)-flowered, 5–6 (–7) × 4–5 (–6) mm. 

Bracteoles ±ovate, brownish, c. 1 mm long; membranous apex fimbriate-ciliate. Tepals 

±equal, usually 2.0–2.3 mm long, ±acuminate to acute, dark brown. Stamens 6; anthers 

0.4–0.5 mm long; filaments 0.5–0.7 mm long; style c. 0.2 mm long; stigmas 0.8–1.0 mm 

long. Capsule ±ovate, with top rounded, ±not mucronate, shorter than or equalling tepals, 

blackish brown to dark castaneous brown; capsule segments ovate, 1.8–2.0 × c. 1.3–1.4 mm. 

Seeds narrowly ellipsoidal, 1.0–1.1 × 0.6 mm, ±pale brown, distally dark apiculate; base 

fibrillate; appendage not visible. 2n=36, Á.Löve & D.Löve, Univ. Colorado Stud. Ser. Biol. 

24: 26 (1966). Fig. 19. 

Northernmost regions of Eurasia and North America, mountains of northern Asia. 10: FIN, 

ICE, NOR, SVA, SWE. 14: RUN. 30: ALT, BRY, CTA, IRK, KRA, TVA, WSB, YAK. 31: 

MAG. 37: MON. 70: ASK, GNL, NWT, YUK. 71: ABT, BRC. 72: QUE, NFL. 75: MAI, NWH, 

VER. Wet, exposed rocky and sandy hillsides, and mountain summits, tundra. Map 79. 

10: NORWAY: Dovre, Knudshoe [Knutshøa], Aug 1875, H.G.Falk s.n. (LD). SVALBARD: Lina Red Bay, 

14 Jul 1899, T.Wulff s.n. (W). SWEDEN: Torne Lappmark, Kiruna, 700 m, G.Simmons 121 (LD). 14. RUSSIA 

NORTH: Lapponia ponojensis, Orlov, A.O.Kihlman 907 (H). 30: WEST SIBERIA: Novaya Zemlya, 

89

90 


Karmakola, 18 Jul 1891, O.J.Ekstam s.n. (LD). KRASNOYARSK: Taymyr, Syndasko R., N.S.Vodop’yanova 

301 (PRA). 70: ALASKA: Monument Ck Road, Teller Road, Williams 1798 (K). GREENLAND: Foulke 

Fjord, Etah, G.Simmons 1481 (LD). NORTHWEST TERRITORIES [Nunavut]: Franklin District, Frobisher 

Bay, Eskimo village, H.A.Senn & J.A.Calder 3941 (LD). 72: QUEBEC: Gaspé, Table-top Mtn, M.L.Fernald 

& J.F.Collins, Pl. East Quebec 195 (LD). 

In some areas of N Europe difficult to distinguish from L. arcuata; in other regions either not 

in contact with the latter, or distinct from it (with occasional, rare hybridization). 

49. Luzula nivalis (Laest.) Spreng., Syst. Veg., 16th edn, 2: 111 (1825) 

Luzula campestris var. nivalis Laest., Kongl. Vetensk.-Akad. Handl. 1822: 334 (1822); Juncus campestris var. 

nivalis (Laest.) Sommerf., Suppl. Fl. Lappon. 16 (1826), n.v.; Juncoides nivalis (Laest.) Coville, Contr. U. S. 

Natl. Herb. 4: 209 (1893). T: Sweden, Lule Lappmark, [lit.: in rupibus occultis Alpium rarissime. In latere 

boreali alpis Jegraapo], norra branten af Jegra-apo-Jalle, 1821, L.L.Laestadius; lecto: S, designated here; 

isolecto: S, UPS [further authentic material: S, UPS, W]. 

Luzula arctica Blytt, Norges Fl. 1: 299 (1861); Luzula arcuata var. hookeriana Trautv., Trudy Imp. 

S.-Peterburgsk. Bot. Sada 1: 80 (1871); Luzula arctica var. typica Buchenau, in H.G.A.Engler, Pflanzenr. 

(iv.36) 25: 69 (1906), nom. inval. T: Norway, Dovre, M.Blytt [Fries, Herb. Norm. Fasc. 10] 65; lecto: UPS, 

designated here. 

Luzula tolmatchewii Kuvaev, Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 99(1): 118 (1994). T: Russia, 

W Siberia, Novaya Zemlya, Insula borealis, inter maris Karskoe [more] et radiostatio Matoczkin Schar, 240 m, 

Tolmatchew 495; holo: LE. 

Illustrations: J.Lid, Norsk & Svensk Fl. 202, fig. 99b (1963); E.Hultén, Fl. Alaska 300 (1968). 

Maps: E.Hultén, Fl. Alaska 300 (1968); E.Hultén & M.Fries, Atlas N. Eur. Vasc. Pl. 1: map 170 (1986), as 

L. arctica; N.K.Kovtonyuk, in L.I.Malyshev & G.A.Peshkova, Fl. Sibiri 4 (Araceae – Orchidaceae): map 36 

(1987) [Siberia]. 

Perennials, usually 8–18 cm tall, caespitose, subglabrous; rhizome ascending; plant base 

straw-brownish or pale castaneous brown. Basal leaves numerous, crowded, ±flat, usually up 

to 5 cm long, (2–) 3–4 mm wide; cauline leaves (1–) 2, 1–2 (–3) cm long, 1–2 mm wide; all 

leaves with obtuse slightly swollen tip with spinuliform mucro (caducous). Inflorescence 

congested in a single dark many-flowered head 0.8–1.0 × 0.6–0.9 cm, composed of 1–3 

clusters; basal cluster sometimes on erecto-patent peduncle up to 3 cm long. Lower bract 

usually scarious to straw-brown, subglabrous, linear-lanceolate, 4–9 mm long, much shorter 

than inflorescence. Bracteoles ±ovate, ±obtuse, sparsely ciliate above, up to 1.2 mm long, 

dark brown. Tepals subequal, 1.6–1.9 (–2.0) mm long, acute, dark castaneous to blackish 

brown. Stamens 6; anthers 0.3–0.6 mm long; filaments 0.4–0.5 mm long; style 0.1–0.2 mm 

long; stigmas 0.8–1.0 mm long. Capsule ovoid-trigonous, exceeding tepals; capsule segments 

2.1–2.3 × c. 1.2 mm, blackish brown. Seeds ellipsoidal, 1.0–1.2 × 0.6–0.7 mm, brown, 

distally with a greyish tip; basal appendage up to 0.2 mm long, fibrillate. 2n=24, fide 

G.Knaben & T.Engelskjøn, Acta Borealia, A. Sci., 21: 4 (1967). 

Arctic regions of Eurasia and North America, mountains of S Siberia. 10: FIN, NOR, SVA, 

SWE. 14: RUN. 30: BRY, CTA, IRK, KRA, TVA, WSB, YAK. 31: MAG. 70: ASK, GNL, 

NWT, YUK. 71: BRC. 72: NFL, QUE. Moist places in arctic and mountain tundra. Map 80. 

10: NORWAY: Folldal, between Råtåsjø and Råtåsjøhø, T.Karlén 211 (LD). SVALBARD: Isfjorden, Green 

Harbour, 5 Jul 1925, Lagerkrantz s.n. (S). SWEDEN: Torne Lappmark, Jukkasjärvi, Mt Nissontjärro, 

1300–1400 m, C.G.Alm & H.Smith 139 (LD); Torne Lappmark, Jukkasjärvi, Torneträsk Lake, K.H.Rechinger 

S–230 (W). 30: WEST SIBERIA: Novaya Zemlya, Krestovaya Guba, 10 Aug 1926, H.Steffen s.n. (LD). 

KRASNOYARSK: E Sayan Mtns, Udinskiy Range, L.I.Malyshev 376 (PRA). 70: GREENLAND: Heilprin 

Land, Brønlundhus, 31 Jul 1963, Fredskild [Pl. Vasc. Groenl. Exs. 268] (UPS); Melville Bugt, Tugtuligssuaq, 

350 m, 16 Aug 1979, Fredskild [Pl. Vasc. Groenl. Exs.] 671 (S). NORTHWEST TERRITORIES [Nunavut]: 

Franklin, Southampton Is., Coral Harbour, O.Hedberg 3315 (UPS); Franklin, Corwallis Is., Resolute, 

O.Hedberg 3248 (UPS).


Presumed hybrids 

Luzula arcuata (Wahlenb.) Sw. subsp. unalaschkensis (Buchenau) Hultén × L. confusa Lindeb. 

Intermediate plants are found at sites with both presumed parents. Up to now known only 

from Alaska [L.Hämet-Ahti, Ann. Bot. Fenn. 10: 128 (1973)]. They usually exhibit a partial 

seed-sterility. 

Luzula arcuata (Wahlenb.) Sw. subsp. arcuata × L. confusa Lindeb. 

Plants intermediate between the two taxa are quite common in Greenland, Scandinavia, Finland 

and W North Russia. The presumed hybrid plants set normal seeds and require further study. 

Unclear Names 

Luzula arcuata f. subnivalis Laest., Bot. Not. 1858: 144 (1858). 

T: Sweden, Qvickjock [Kvikkjokk], L.L.Laestadius; syn: n.v. 

Luzula arcuata f. latifolia Laest., Bot. Not. 1858: 144 (1858). 

T: Sweden, Åreskutan, 1819, L.L.Laestadius; syn: n.v. 

Luzula hyperborea var. minor Hook., Fl. Bor. Amer. 2: 188 (1840) 

Juncoides hyperborea var. minor (Hook.) E.Sheld., Bull. Geol. Nat. Hist. Surv. 9: 64 (1894), n.v. T: n.v. 

Sect. 7. LUZULA 

Luzula sect. Gymnodes Griseb., Spic. Fl. Rumel. 2: 405 (1846); Gymnodes (Griseb.) Fourr., Ann. Soc. Linn. 

Lyon, sér. 2, 17: 172 (1869); Luzula subg. Gymnodes (Griseb.) Buchenau, Bot. Jahrb. Syst. 1: 114 (1880); 

Luzula sect. Apterodes Rouy, Fl. France 13: 256 (1912), nom. illeg. T: not designated. 

Luzula subsect. Ulea Novikov, Byull. Moskovsk. Obshch. Isp. Prir., Odt. Biol. 95(6): 68 (1990). T: Luzula 

ulei Buchenau 

Luzula subsect. Caricinae Novikov, Byull. Moskovsk. Obshch. Isp. Prir., Odt. Biol. 95(6): 68 (1990). 

T: Luzula caricina E.Mey. 

Luzula subsect. Pumilae Novikov, Byull. Moskovsk. Obshch. Isp. Prir., Odt. Biol. 95(6): 69 (1990). 

T: Luzula pumila Hook.f. 

Luzula subsect. Sibiricae Novikov, Byull. Moskovsk. Obshch. Isp. Prir., Odt. Biol. 95(6): 69 (1990). 

T: Luzula sibirica (V.I.Krecz.) V.I.Krecz. 

Luzula ser. Inaequales K.F.Wu, J. E. China Norm. Univ., Nat. Sci. Ed., 1992(3): 97 (1992) 

Perennials. Leaves with obtuse, usually slightly swollen tips. Lower bract herbaceous. 

Inflorescence wholly congested or umbelloid, decompound, of one to numerous spike- or 

head-like clusters each usually 3–many-flowered. Seed appendage basal, usually conspicuous, 

0.1–0.8 mm long, fibrillate or not so. Diploids, true polyploids and complete or partial 

agmatoploids. 

A section of 57 species with several diversity centres in Europe, the Far East, Australia, New 

Zealand, W North America and S America, grouped in this treatment by their regions of 

predominant occurence. There is a very low level of major structural differentiation within 

the section, and many taxa are difficult to distinguish from one another. In some regions 

(W North America, China) the section is in need of taxonomic revision. 

Successful identification by means of the following key requires careful measurement of 

well developed material, preferably of more than a single plant. Method of measurement, as 

given in the introductory part to the genus, should be observed. 

91

92 


1 Stamens 3 74. L. leptophylla 

1: Stamens 6 

2 Most flowers in an inflorescence borne singly, some in few-flowered 

spike-like clusters 102. L. hawaiiensis 

2: Most flowers in pedunculate clusters, or inflorescence wholly congested 

3 Plants in dense compact cushions 

4 Flowering stems abbreviated, hidden in leaf rosettes, not elongating 

as fruits mature; seeds broadly ovoid, 0.7–0.9 mm wide 79. L. colensoi 

4: Flowering stems at least slightly overtopping leaves, elongation 

during fruit ripening; seeds ±ellipsoid, 0.5–0.6 mm wide 

5 Leaves inserted (imbricate) along proximal 2/3– 3/4 of stem, 

0.4–0.7 mm wide; inflorescence 1–5-flowered 76. L. crenulata 

5: Leaves mostly basal to subbasal, 0.7–1.3 mm wide; inflorescence 

4–10 (–20)-flowered 78. L. pumila 

3: Plants caespitose to loosely caespitose, with or without creeping 

rhizomes, sometimes forming mats but never dense compact cushions 

6 Capsule top or upper part mostly densely papillose 

7 Densely caespitose; rhizome short or oblique-ascending; stolons absent 

8 Seed appendage 0.4–0.6 mm 

9 Tepals with margins indistinct or absent 83. L. meridionalis 

9: Tepals with distinct broad whitish-scarious margins 84. L. flaccida 

8: Seed appendage 0.3 mm or less 

10 Leaf margin usually smooth, rarely sparsely papillose-serrulate 

10: Leaf margin densely to sparsely papillose-serrulate 

88. L. novae-cambriae 

11 Basal leaves usually 4–7 (–10) mm wide 

11: Basal leaves up to 3.5 mm wide 

86. L. australasica 

12 Seeds 0.5–0.6 mm wide; appendage c. 0.1 mm long 90. L. alpestris 

12: Seeds c. 0.8 mm wide; appendage 0.2–0.3 mm long 93. L. poimena 

7: Loosely caespitose; rhizomatous plants with stolons 

13 Style abbreviated, to 0.2 mm long; inflorescence a single dark 

head 0.7–1.0 cm in diam. 

13: Style 0.3–0.5 mm long; inflorescence composed of (1–) 2–4 (–5) 

subsessile subcylindrical clusters each 5–12-flowered and c. 6–9 mm 

long; proximal cluster often ±remote; other clusters approximated, 

92. L. atrata 

forming elongated lobulate head 1–2 (–3) cm long 89. L. modesta 

6: Capsule smooth in most flowers, rarely with sparse papillae 

14 Peduncles ±densely papillose in the middle part (under 

magnification of at least 80×) 

15 Tepals ±equal; seeds 0.7–0.8 mm wide; clusters variable, often 

less than 10-flowered 

16 Clusters (5–) 6–11 (–13), usually 8–11-flowered; peduncles 

frequently with secondary branches; style 0.3–0.4 mm long 

16: Clusters 3–5 (–7), usually 3–7-flowered; peduncles without 

59. L. multiflora 

secondary branches; style 0.7–1.0 mm long 53. L. fallax 

15: Outer tepals conspicuously longer; seeds 0.5–0.6 mm wide; 

clusters usually more than 10-flowered 50. L. pallescens


14: Peduncles smooth in the middle part 

17 Flower clusters loose at base and/or some flowers ±remote 

18 Capsule segments 1.9–2.1 mm long; seeds 0.6–0.7 mm wide 

[S Italy] 52. L. calabra 

18: Capsule segments longer than 2.3 mm, seeds usually wider than 

0.7 mm [NW and C America] 

19 Style 0.6–1.2 mm long; seeds 0.9–1.1 mm wide 98. L. subsessilis 

19: Style up to 0.6 mm long; seeds 0.7–0.9 mm wide 

20 Tepals dark to blackish brown, 2.0–2.8 mm long 103. L. caricina 

20: Tepals pale straw-brown to pale brown, (2.5–) 3.0–4.0 mm long 99. L. comosa 

17: Flower clusters (evenly) dense, without remote flowers 

21 Seed appendage usually 0.5 mm or more 

22 Inflorescence wholly congested, or sometimes some of the 

clusters on erect peduncles 

23 Anther/filament length ratio c. 1.0; basal leaves usually 

3.5–6.0 mm wide 58. L. congesta 

23: Anther/filament length ratio c. 2.0; basal leaves usually 

2.0–3.5 mm wide 68. L. capitata 

22: Inflorescence of pedunculate clusters; if partly congested then 

some of the peduncles flexuose or recurved and nodding 

24 Plants with long creeping rhizomes and frequent ascending 

stolons 56. L. campestris 

24: Plants densely to loosely caespitose; rhizomes short or shortascending; 

stolons absent 

25 Rhizome short-ascending, short-noded, usually with a few 

pale swollen reduced leaves; flower clusters cylindric to 

±ovoid 101. L. bulbosa 

25: Rhizome short, vertical, without swollen reduced leaves; 

flower clusters broadly ovoid to hemispherical 

26 Capsule distinctly exceeding perianth 71. L. lutescens 

26: Capsule shorter than perianth 

27 Seeds 0.9–1.0 mm wide; anthers usually longer than 

2.0 mm [C and SC Europe] 55. L. divulgata 

27: Seeds 0.6–0.8 mm wide; anthers up to 1.5 mm long 

[E North America] 100. L. echinata 

21: Seed appendage usually 0.4 mm or less (very rarely to 0.5 mm) 

28 Leaf margins ±smooth, without dense papillose serrulation, 

rarely with solitary, often reddish papillae 

29 Leaf tip acute to subobtuse with an acute mucro 91. L. acutifolia 

29: Leaf tip obtuse, often ±swollen 

30 Rhizome creeping-ascending, branched, plants therefore 

very loosely caespitose; stolons present or absent 

31 Stem reduced, to 2 cm long; inflorescence usually 2–10flowered 

80. L. decipiens 

31: Stem longer than (10–) 15 cm; inflorescence usually 

many-flowered 

93

94 


32 Inflorescence usually of 1–2 (–4) elongated heads 

c. 5–20 × 5–10 mm, each composed of 3–7 (–10) spikelike, 

subcylindrical, shortly pedunculate to subsessile 

clusters [Papua-New Guinea] 72. L. papuana 

32: Inflorescence ±umbelloid, of one to several subsessile 

and several pedunculate clusters [New Zealand] 81. L. picta 

30: Rhizome short, indistinct or ascending, unbranched, plants 

therefore densely caespitose; stolons absent 

33 All but 1–2 (–3) clusters pedunculate 

34 Style 0.5–0.9 mm long; stigmas 1.5–2.5 mm long 

[Hawaiian Is.] 102. L. hawaiiensis 

34: Style usually 0.9–1.5 mm long; stigmas 2.0–3.5 mm 

long [New Zealand] 82. L. banksiana 

33: All or most clusters congested, forming a single head; 

occasionally, some clusters may be pedunculate 

35 Seed appendage 0.2–0.3 mm long 87. L. ovata 

35: Seed appendage 0.1 (–0.2) mm long 

36 Tepals dark brown to blackish; margins dark 

castaneous, narrow; seeds 0.5–0.6 mm wide 75. L. crinita 

36: Tepals pale brown to deep castaneous-brown; 

margins distinct, broad, pale brownish to scarious; 

seeds 0.6–0.8 mm wide 

37 Tepals usually 1.7–2.5 mm long 77. L. rufa 

37: Tepals usually 2.6–4.0 mm long 

38 Basal leaves usually 3–8 mm wide 82. L. banksiana 

38: Basal leaves usually 1.5–4 mm wide 

39 Basal leaves stiff, canaliculate, with margins subglabrous 

to sparsely ciliate; seeds 0.6–0.7 mm 

wide [New South Wales] 86. L. australasica 

39: Basal leaves ±flaccid, not distinctly stiff, 

with margins densely to sparsely ciliate; seeds 

c. 0.8 mm wide [Chile] 104. L. tristachya 

28: Leaf margin ±densely papillose-serrulate 

40 Capsule distinctly exceeding perianth; tepals to 1.8 mm long 69. L. oligantha 

40: Capsule as long as perianth or shorter; tepals longer than 1.8 mm 

41 Inflorescence of 1–5 (–6)–flowered clusters; clusters 

numerous, more than 15 [Hawaiian Is.] 102. L. hawaiiensis 

41: Inflorescence of (5–) 6–many-flowered clusters; clusters 

few, rarely to 15 

42 Anther/filament length ratio 3–5 73. L. philippinensis 

42: Anther/filament length ratio 0.5–3 

43 Tepals usually 4.5–6.0 mm long [Lord Howe Is.] 94. L. longiflora 

43: Tepals usually up to 4.2 mm long 

44 Seeds usually 0.9–1.0 mm wide; seed appendage 

0.4–0.6 mm long; seeds 1.2–1.5 mm long; tepals 

straw-brown to mid-brown 58. L. congesta 

44: Seeds up to 0.8 mm wide; seed appendage to 0.4 mm 

long; seeds 0.8–1.2 mm long, if seeds up to 1.3 mm 

then tepals dark castaneous-brown to blackish-brown


45 Plant base usually bulbous; seed appendage usually 

0.3–0.4 mm long; seeds 0.8–0.9 mm long [Australia] 85. L. densiflora 

45: Plant base not bulbous; seed appendage 0.1–0.3 mm 

long; if appendages 0.3–0.4 mm long then seeds 

1.0 mm or longer 

46 Tepals up to 2.4–2.5 mm long 

47 Seeds 0.5–0.6 mm wide 

48 Plants with long, branched, oblique or ascending 

rhizomes; stolons present or absent 

49 Anthers c. twice as long as filaments; seed 

appendage c. 0.2 mm long 65. L. abchasica 

49: Anthers ± as long as filaments; seed appendage 

to 0.1 mm long 57. L. sudetica 

48: Plants densely caespitose; rhizome short; stolons 

absent 

50 Style 0.4–0.6 mm long [Caucasus and NE 

Anatolia] 64. L. stenophylla 

50: Style 0.1–0.3 mm long 

51 Inflorescence with 1–3 pedunculate clusters 

(in addition to subsessile ones); peduncles 

1.5–4.0 cm long [Japan] 71. L. nipponica 

51: Inflorescence wholly congested, or rarely one 

cluster on peduncle to 1.5 cm long [North 

America or Greenland] 

52: Tepals dark, usually blackish brown; 

seed appendage c. 0.2 mm long; seeds 

0.8–0.9 (–1.0) mm long [California] 97. L. orestera 

52: Tepals brown to castaneous-brown; 

seed appendage c. 0.1 mm long; seed 

1.0–1.1 mm long [arctic and subarctic North 

America and Greenland] 96. L. groenlandica 

47: Seeds 0.7–0.8 mm wide 

53 Plants with long, branched stoloniferous 

rhizomes [Tropical Africa] 60. L. mannii 

53: Plants with short rhizomes; stolons absent 

[outside Africa] 

54 Pedunculate clusters 5–30; peduncles with 

numerous secondary branches; clusters 2–7flowered; 

leaves 4.5–11 mm wide [South 

America] 106. L. ulei 

54: Pedunculate clusters 1–3; secondary branches 

usually absent; clusters more than 5-flowered; 

leaves to 3.5 mm wide [outside South America] 

55 Inflorescence with 1–3 pedunculate clusters 71. L. nipponica 

55: Inflorescence usually wholly contracted (or 

basal cluster remote) 

56 Seed appendage 0.2–0.3 mm long 87. L. ovata 

56: Seed appendage 0.1 mm long 

57 Lower bract tip acute-mucronate to acute; 

basal cluster usually remote [Taiwan] 67. L. taiwaniana 

95

96 


57: Lower bract tip obtuse, ±swollen; basal 

cluster is a part of the wholly congested 

inflorescence [North America or 

Greenland] 96. L. groenlandica 

46: Tepals longer than 2.6 mm 

58 Seed appendage c. 0.1 mm long 

59 Outer tepals distinctly aristate [S India] 63. L. indica 

59: Outer tepals usually acute to acuminate, not 

aristate [outside India] 

60 Anther/filament length ratio 2.0 (–3.0) [South 

America] 105. L. ostenii 

60: Anther/filament length ratio up to 1.5 

61 Seeds 0.5–0.6 mm wide 

62 Style 0.1–0.3 mm long 

63 Plants with long branched ascending 

rhizomes; tepals with indistinct narrow 

margins; tepals often unequal (outer 

longer) [Europe] 57. L. sudetica 

63: Plants densely caespitose; rhizome short, 

unbranched; tepals with distinct broad 

margins; tepals ±equal [California] 97. L. orestera 

62: Style 0.4–0.6 mm long 

64 Inflorescence almost completely contracted, 

elongated, 2–5 cm long, often interrupted at 

base; basal leaves usually 5–6 mm wide; 

tepals castaneous-brown [E Africa] 61. L. abyssinica 

64: Inflorescence contracted, subglobose, to 

1.8 cm long, compact; basal leaves usually to 

3–4 mm wide; tepals dark to blackish brown 

[Caucasus and NE Anatolia] 64. L. stenophylla 

61: Seeds 0.7–0.8 mm wide 

65 Inflorescence ±wholly contracted, 

sometimes with 1–3 pedunculate clusters; 

peduncles without secondary branches 

66 Style 0.4–0.6 mm long; inflorescence 

elongated, 2–5 cm long [E Africa] 61. L. abyssinica 

66: Style 0.3–0.4 mm long, congested part of 

inflorescence to 1.5 cm long [China and 

Tibet] 66. L. bomiensis 

65: Inflorescence of 1–4 subsessile and 3–many 

pedunculate clusters (if subsessile clusters 3 

or 4 then peduncles with secondary 

branches) 

67 Style 0.4–0.5 mm long; tepals 2.3–3.0 mm 

long [Tropical Africa] 60. L. mannii 

67: Style 0.5–0.9 mm long; tepals 2.8–4.4 mm 

long [Hawaiian Is.] 102. L. hawaiiensis 

58: Seed appendage 0.2 mm or longer 

68 Seeds 0.5–0.6 mm wide


69 Plants with creeping-ascending branched 

rhizomes and stolons 65. L. abchasica 

69: Plants densely caespitose; rhizome short; 

stolons absent 97. L. orestera 

68: Seeds usually 0.7–0.8 mm wide 

70 Cauline leaves conspicuously broad, 

(4.5–) 5.0–8.0 (–12.0) mm wide 

71 Seed appendage 0.1–0.2 mm long [Hawaiian Is.] 102. L. hawaiiensis 

71: Seed appendage 0.2–0.4 mm long [Beringian 

region] 95. L. kobayasii 

70: Cauline leaves not conspicuously broad, 

usually to 4 mm wide 

72 Inflorescence wholly or almost completely 

congested 

73 Style 0.3–0.4 mm long [China and Tibet] 66. L. bomiensis 

73: Style 0.6 mm or more long 

74 Capsule segments usually 2.2–2.4 mm 

long; tepals straw-brown to brown; seeds 

0.7–1.0 mm long [South Africa] 62. L. africana 

74: Capsule segments usually 2.5–2.8 mm 

long; tepals dark brown; seeds 1.0–1.2 mm 

long [European Alps] 51. L. alpina 

72: Inflorescence of mostly pedunculate clusters 

75 Style usually 0.8–1.0 mm long; 

anther/filament length ratio usually 

(1.5–) 2.0 (–3.0) 

76 Seed appendage 0.1–0.2 mm long [South 

America] 105. L. ostenii 

76: Seed appendage usually 0.3–0.4 mm long 

[S Europe] 54. L. taurica 

75: Style usually 0.3–0.8 mm long; 

anther/filament length ratio usually 1.0–2.0 59. L. multiflora 

The species of this large Section are grouped below, for convenience, by their regions of 

predominant occurence. 

PREDOMINANTLY EUROPEAN SPECIES 

50. Luzula pallescens Sw., Summa Veg. Scand. 13 (1814) 

Juncus campestris var. lapponicus Retz., Fl. Scand. Prodr., 2nd edn, 82 (1795); Juncus pallescens Wahlenb., 

Fl. Lapp. 87 (1812), nom. illeg., non Lam. (1789); Luzula campestris subvar. pallescens (Sw.) E.Mey., Syn. 

Luzul. 18 (1823); Luzula campestris var. pallescens (Sw.) Wahlenb., Fl. Suec. 1: 218 (1824), nom. illeg.; 

Luzula multiflora var. pallescens (Sw.) W.D.J.Koch, Syn. Fl. Germ. Helv., 2nd edn 2, 847 (1844); Luzula 

erecta var. pallescens (Sw.) Nolte ex Sond., Fl. Hamburg. 200 (1851); Luzula campestris subsp. pulchella 

Čelak., Prodr. Fl. Böhmen 1: 85 (1867); Luzula erecta subsp. pallescens (Sw.) Nym., Consp. Fl. Eur. 752 

(1882); Luzula sudetica [Rasse] pallescens (Sw.) Asch. & Graebn., Syn. Fl. Mitteleur. 2(2): 519 (1904); 

Luzula campestris subsp. pallescens (Sw.) Čelak., Prodr. Fl. Böhmen 4: 749 (1881); 

97

98 


Juncoides pallescens (Sw.) Druce, Bot. Soc. Exch. Club Brit. Isles 7: 899 (1926); Luzula multiflora subsp. 

pallescens (Sw.) Reichg., Fl. Neerl. 1: 208 (1964). T: [Sweden, Lappland]; lecto: LINN449.47, designated 

here. 

Luzula erecta var. micrantha Schltdl., Fl. Berol. 1: 204 (1823). T: Germany, [Berlin], D.F.L.Schlechtendal. 

syn: B, destroyed; syn: HAL69172. 

Luzula pallescens var. umbrosa Čelak., Oesterr. Bot. Z. 11: 312 (1861). T: [Czech Republic] Johannisbad 

[Jánské Lázně], J.Kablik; lecto: PR, designated here. 

Luzula multiflora var. minor Bluff & Fingerh., Comp. Fl. German. 1: 444 (1825). T: [Germany, otherwise 

not indicated] 

Luzula pallescens var. umbrosa V.I.Krecz., Žurn. Russk. Bot. Obshch. Akad. Nauk S.S.S.R. 12: 491 (1928). 

T: not designated. 

Luzula pallescens var. communis V.I.Krecz., Žurn. Russk. Bot. Obshch. Akad. Nauk S.S.S.R. 12: 491 (1928). 


Luzula pallescens var. arenaria V.I.Krecz., Žurn. Russk. Bot. Obshch. Akad. Nauk S.S.S.R. 12: 491 (1928). 


Luzula pallescens var. ruderalis V.I.Krecz., Žurn. Russk. Bot. Obshch. Akad. Nauk S.S.S.R. 12: 491 (1928). 


Luzula pallidula Kirschner, Taxon 39: 110 (1990). T: Czech Republic, Moravia, Hodonín, in silva Dúbrava, 

10 Jul 1987, R.Řepka & J.Kirschner; holo: PR. 

Illustrations: T.Szynal & J.Mądalski, Atlas Fl. Polsk. 2(2): 147 (1931); J.Kirschner, Folia Geobot. Phytotax. 

28: 168, fig. 8 (1993). 


N. Eur. Vasc. Pl. 1: map 176 (1986); N.K.Kovtonyuk, in L.I.Malyshev & G.A.Peshkova, Fl. Sibiri 4 (Araceae 

– Orchidaceae): map 37 (1987) [Siberia]. 

Perennials, 9–15 (–30) cm tall, densely caespitose; without stolons; rhizome short; stems erect. 

Basal leaves densely to subsparsely ciliate, 2.0–3.5 mm wide; cauline leaves usually 2 or 3, 

with densely papillose-serrulate margin, upper cauline leaves 3.0–5.5 cm long, 2.0–3.0 mm 

wide. Lower bract usually 3.0–4.0 cm long, overtopping or rarely shorter than inflorescence. 

Inflorescence composed of 1–2 subsessile and (5–) 7–12 (–18) pedunculate clusters; clusters 

±cylindrical, 4–7 × 3–4 mm long, usually (9–) 11–14 (–20)-flowered. Peduncles straight, 

densely papillose, often with secondary branches, with longest peduncles (including clusters) 

2.0–5.5 cm long. Bracteoles 0.8–1.2 mm long, ciliate. Tepals pale greenish to membranous to 

straw-brown; outer tepals conspicuously longer than inner, 2.0–2.8 mm long, subacuminate. 

Anthers 0.5–0.6 mm long, ±as long as c. 0.5 mm long filaments; ovary 0.5–0.6 mm long; 

style 0.2–0.3 mm long; stigma early deciduous, 0.6–0.7 mm long. Capsules obovoid to 

subglobular, equalling or shorter than perianth; capsule segments 1.6–1.9 mm long. Seeds 

ovoid, 0.7–0.8 mm long, 0.5–0.6 mm wide; appendage 0.2–0.3 mm long. 2n=12 [12AL], fide 

J.Kirschner, Thaiszia 2: 28–29 (1992). Fig. 21. 

Widely distributed in N Eurasia, from Great Britain and Scandinavia to the Russian Far East 

and China. 10: DEN, FIN, GRB, NOR, SWE. 11: AUT, CZE, GER, HUN, POL. 13: ROM, 

YUC. 14: BLR, BLT, KRY, RUC, RUE, RUN, RUS, RUW, UKR. 30: ALT, BRY, CTA, 

IRK, KRA, TVA, WSB, YAK. 31: AMU, KAM, KHA, KUR, MAG, PRM, SAK. 32: KAZ, 

KGZ, TKM, TZK, UZB. 33: NCS, TCS. 36: CHC, CHI, CHM, CHN, CHX. 37: MON. 

38: JAP, KOR. 40: JMK. 72: nfl, que. Wood clearings and tracks, and disturbed meadows on 

sandy or gravelly substrates. Map 81. 

10: NORWAY: Hallingdal, Nesbyen, G.Samuelsson 401 (UPS). SWEDEN: Sala, Heby, Huddunge, 

J.Kirschner L300 (PRA). 11: CZECH REPUBLIC: N Moravia, Vidnava, J.Hruby [in F.Petrak, Fl. Bohem. 

Morav. Exs. 513] (LD). 13: BOSNIA: Fojnica, 600 m, 18 Sep 1889, S.Murbeck (LD). 14: MOLDOVA: 

Iaşi, Uricani, C.Dobrescu & E.Eftimie [Fl. Mold. Dobrog. Exs. 189] (PR). 30: WEST SIBERIA: 

Novosibirsk, Klyuchi, D. Shaulo, cult. no. JK154 (PRA). 31: RUSSIA: Kamchatka, Petropavlovsk, 

E.Hultén 192 (S). 32: KIRGIZISTAN: Tian-Shan, Kirgizski Range, NE of Pishpek, valley of R. Ala Archa, 

1700–200 m, 28 May 1974, V.Vašák (W). 33: NORTH CAUCASUS: Mt Elbrus region, Adyrsu Valley, 

1600–2000 m, 24 Jun 1980, V.Vašák (W). 36: CHINA: Jilin, Wo-chin-kow, O-muh Hsien, 800 m, 

H.W.Kung 1891 (S). 37. MONGOLIA: E Mongolia, W part of Great Khingan, 19 Aug 1973, N.Dorofeyuk 

(MW). 38: KOREA: Cham-gion Province, Tumin-gan R., V.L.Komarov 356 (BM). 40: WEST 

HIMALAYA, KASHMIR: Gulmarg, 1600–1700 m, 16 Apr 1948, J.J.Thomson (K). 72: QUEBEC: 

Bonaventura Co., Giroux Station, Maria, 11 Jul 1905, Williams, J.F.Collins & M.L.Fernald (UPS).


Figure 21. Luzula pallescens. A, habit; B, flower cluster on papillose peduncle; C, flower; 

D, capsule; E, seed; F, G, tepals; H, stem T.S.; I, leaf T.S.; J, detail of peduncle. 

(A–J, B.Błocki, 9 Jun 1891, KRA). Scale bars: A = 5 cm; B = 5 mm; C–G = 2 mm; 

H, I = 1 mm; J = 0.5 mm. Drawn by T.Szynal. Reproduced with permission, from T.Szynal 

& J.Mądalski, Atlas Fl. Pol. 2/2: 147 (1931). 

99

100 


Characterized by numerous small flowers with unequal tepals, and densely papillose peduncles 

(under the microscope). Sometimes confused with pale-flowered forms of L. multiflora. 

51. Luzula alpina Hoppe, in J.W.Sturm, Deutschl. Fl. 18 (77): no. 6 (1839) 

Luzula multiflora var. alpina (Hoppe) Willk., Führer deut. Pfl., 2nd edn, 282 (1882); Luzula sudetica proles 

alpina (Hoppe) Asch. & Graebn., Syn. Mitteleur. Fl. 2(2): 518 (1904). T: Austria, Heiligenblut, D.H.Hoppe 

[Herb. Viv. Pl. Gram.] 108; lecto: BRNM, fide J.Kirschner, T.Engelskjøn & G.S.Knaben, Preslia 60: 98 

(1988); isolecto: LI, HAL. 

Luzula campestris β latifolia Gaudin, Fl. Helv. 2: 572 (1828). T: [Swiss Alps] in M. Gemmii, N.C.Seringe; 

holo: n.v. 

Luzula multiflora var. alpestris Beyer, Verh. Bot. Vereins Prov. Brandenburg. 41: xxii (1900); Luzula 

campestris var. alpestris (Beyer) Vollm., Fl. Bayern 138 (1914), nom. illeg., non G.Mey. (1836); Luzula 

campestris subsp. alpestris (Beyer) Kožuharov, Izv. Bot. Inst. (Sofia) 11: 132 (1963), nom. illeg., non G.Mey. 

(1836) T: [numerous syntypes from the Alps, mostly B, destroyed]. 

Luzula campestris var. vallesiaca Beauverd, Bull. Soc. Bot. Genève, ser. 2, 10: 285 (1918). T: Switzerland, 

Valais, Zeneggen, c. 1330 m, G.Beauverd; holo: G. 

Illustrations: J.Kirschner, T.Engelskjøn & G.S.Knaben, Preslia 60: 99, fig. 2, Plate 1, 2, 3 (1988); J.Kirschner, 

Folia Geobot. Phytotax. 28: 149, fig. 1 (1993). 

Map: J.Kirschner, T.Engelskjøn & G.S.Knaben, Preslia 60: 104, fig. 4 (1988). 

Perennials, (8–) 10–20 (35) cm tall, densely caespitose, without stolons, sparsely ciliate; 

stems erect, ±stout. Upper cauline leaves 4–5 cm long, 3–4 mm wide; basal leaves 3–5(–7) mm 

wide. Leaf margin remotely papillose-serrulate. Basal bract usually spreading, overtopping 

inflorescence. Inflorescence congested, composed of (3–) 5–6 (–8) sessile to subsessile 

(rarely some pedunculate) clusters; peduncles smooth; clusters usually of 7–10 flowers. Tepals 

±lanceolate, gradually tapering to long narrow sharp pointed tips, usually dark brown; outer 

tepals (2.7–) 3.0–3.5 (–3.7) mm long, ±equalling the inner ones. Anthers (0.8–) 0.9–1.1 mm 

long, at most 1.5 times longer than the 0.5–0.8 mm long filaments. Style c. 0.6 mm long, 

shorter than the ovary. Capsule segments usually 2.5–2.8 mm long, shorter than the perianth, 

±obovate. Seeds narrowly ovoid, 1.0–1.2 mm long, 0.7–0.8 mm wide; appendages 0.3–0.4 mm 

long. 2n=36 [12AL + 24BL], fide J.Kirschner, T.Engelskjøn & G.S.Knaben, Preslia 60: 

100–101 (1988). 

Endemic to the Alps and E Pyrenees, from Switzerland, Italy and Slovenia northwards to 

S Germany (Bavaria). 11: AUT, GER, SWI. 12: FRA, SPA. 13: ITA, YUC-SL. Map 82. 

11: AUSTRIA: Lechtal, Elpigenalp, Bernhardseck, 2200 m, O.Sebald 8235 (M); Carinthia, Greifenburg, 

Embergeralm, 1790 m, 21 Jul 1949, Eggler (GZU); Carinthia, Gailtaler Alpen, N of St. Jakob, ‘Auf der Mussen’, 

19 Jul 1998, L.Kirschnerová (PRA); Tirol, Stubaital, D.Podlech [Soc. Éch. Pl. Vasc. Eur. Occid, no 8876] (G). 

GERMANY: Bavaria, Berchtesgaden, Mittereis-Alm, W.Lippert 25854 & D.Podlech (M); Oberstdorf, 

Fallhorn, 1650 m, 10 Jul 1923, Hirth (W). SWITZERLAND: St. Gallen, Vältis, Brändlisberg, 1908, 

H.Schinz (Z). 12: SPAIN: Huesca, Torla, Ordesa, Gradas Soaso, 1850 m, P.Montserrat (JACA 478371). 

ANDORRA: Coll Blanch d’Envalira, 2500 m, 18 May 1948, T.M.Losa (BCF 4077). 13: ITALY: Dolomiti, 

Vigo di Fassa, 30 Jul 1956, E.Werdermann & Meyer (K). 

Close to Luzula multiflora but usually with a congested dark inflorescence and broad basal 

leaves. In the Alps, L. alpina is the only member of the genus with the peculiar mixed 

karyotype. 

52. Luzula calabra Ten., Fl. Napol. 3: 386 (1829) 

Luzula multiflora var. calabra (Ten.) Parl., Fl. Ital. 2: 307 (1852); Luzula multiflora subsp. calabra (Ten.) 

Arcang., Comp. Fl. Ital. 713 (1882); Luzula campestris var. calabra (Ten.) Buchenau, Bot. Jahrb. Syst. 12: 

163 (1890). T: [Italy, Calabria] Paludi della Sila, Thomas; lecto: NAP, fide J.Kirschner, Folia Geobot. 

Phytotax. 28: 148 (1993). 

Illustrations: Miggliaccio, Giorn. Bot. Ital. 71: 334, fig. 1, 2 (1964); S.Pignatti, Fl. Ital. 3: 446 (1982); 

J.Kirschner, Folia Geobot. Phytotax. 28: 151, fig. 2 (1993). 

Perennials, 25–45 cm tall, densely caespitose, without stolons; rhizome short; stems erect. 

Basal leaves densely to subsparsely ciliate, 1.7–2.2 mm wide; cauline leaves usually 1 or 2, 

with densely papillose-serrate margins; upper cauline leaves 5–7 cm long, 2.0–2.8 mm wide.


Lower bract usually 2.0–4.0 cm long, shorter than inflorescence. Inflorescence with one 

subsessile and 1–6 pedunculate clusters; peduncles straight, smooth, usually without 

secondary branches; longest peduncles (including clusters) 2.0–9.5 cm long. Clusters 

±cylindrical, 8–25-flowered; basal flowers of the pedunculate clusters often slightly remote. 

Bracteoles 1.6–1.9 mm long, ciliate or fimbriate-ciliate. Tepals pale brown; outer tepals 

±equalling or slightly longer than inner ones, 2.4–2.8 mm long, subabruptly narrowing at the 

apex. Anthers 0.8–0.9 mm long; filaments c. 0.4 mm long; ovary 0.7–0.8 mm long; style 

0.3–0.4 mm long; stigma early deciduous, 0.8–1.0 mm long. Capsules subacuminate, 

equalling or shorter than perianth; capsule segments 1.9–2.1 mm long. Seeds oblong-ovoid, 

c. 0.9 mm long, 0.6–0.7 mm wide; appendage 0.2–0.3 mm long. 2n=24 [probably 24BL], fide 

Virzo & Miggliaccio, Delpinoa 6–7: 81 (1965). 

Endemic to S Italy. 13: ITA. Grassy slopes, stream banks, meadows. Map 83. 

13: ITALY: Calabria, Magna Sila, Botte Donato, c. 1800 m, 16 Aug 1909, Cavara & Grande 28 (BM, P); 

Catanzaro, Sila Piccola, from Taverna Villagio Mancusa to Butoro, c. 1200 m, 16 Jun 1979, P.H.Davis & 

D.A.Sutton (BM); Sila, W of Lorica, Sila, Mt Sorbello, c. 1850 m, 18 Jun 1979, P.H.Davis & D.A.Sutton 

(BM); Calabria, E ridge of Montenero, 1400–1600 m, S.Snogerup & B.Snogerup 2081 (LD). 

Rare endemic distinct in having slightly loose clusters at the base of the inflorescence; in 

general habit it is similar to L. pallescens. 

53. Luzula fallax Kirschner, Preslia 63: 84 (1991) 

T: N Greece, Mt. Pangeon, 1800–1850 m. S.Snogerup & A.Strid 4934, cult. no. L134; holo: PR, iso: LD, PR. 

Illustration: J.Kirschner, Preslia 63: 85, fig. 1 (1991). 

Perennials usually 15–20 cm tall, densely caespitose; stolons absent; rhizome short; stems erect, 

slender. Basal leaves densely to subsparsely ciliate, 2–4 mm wide. Cauline leaves usually 2, 

with densely papillose-serrate margin; upper cauline leaves 1.5–4.5 cm long, 2.0–3.0 mm 

wide. Lower bract usually 1.2–2.5 cm long, shorter than inflorescence. Inflorescence 

composed of 1–2 subsessile and 2–5 pedunculate clusters; peduncles straight or flexuose, 

papillose in upper half or papillose throughout, usually without secondary branches; longest 

peduncles (including clusters) 2.0–5.5 cm long. Clusters ovoid to hemispherical, 4.5–7.5 × 

5.0–6.5 mm, usually (2–) 4–6 (–8)-flowered; basal flowers of the pedunculate clusters not 

remote. Bracteoles 1.0–1.5 mm long, ciliate above. Tepals pale to dark brown; outer tepals 

±equalling inner ones, 2.6–3.0 mm long. Stamens 6; anthers (0.5–) 0.7–0.9 mm long; 

filaments 0.4–0.6 mm long; ovary 0.7–0.8 mm long; style 0.7–1.0 mm long; stigma 1.7–2.2 mm 

long. Capsules obovoid to subglobular, equalling or shorter than perianth; capsule segments 

brown, 2.0–2.4 mm long, c. 1.3 mm wide. Seeds ovoid, (0.9–)1.0 mm long, 0.7–0.8 mm 

wide; appendage c. 0.3 mm long. 2n=24 [24BL], fide J.Kirschner, Preslia 63: 86 (1991). 

Balkan Peninsula from Bulgaria to European Turkey. 13: ALB, BUL, GRC, TUE. Slopes of 

hills and lower mountains and meadows on naturally or artificially disturbed acid soils. 

Map 84. 

13: ALBANIA: Kiri near Skutari, I.Dörfler 43 (LD). BULGARIA: Pirin [Mtns], Melnik, slopes above 

Rozhen, Vlashkia Put, J.Kirschner & B.Kuzmanov L116 (PRA); Sofia, Lyulin Planina, 22 Jun 1919, Akhtaroff 

(SOM). GREECE: Mt Pangeon, 0.5 km E of summit, S.Snogerup & A.Strid 4934 (LD). Pierias, NW 

foothills of Mt Olympos, A.Strid & Andersen 8440 (LD); Phocis, Mt Giona, c. 2000 m, 

L.-Å.Gustavsson 6909 (LD); Trikala, SW of Pirra, Mt Augo, B.G.Aldén 2312 (LD); Evritania, Gardikion, 

Mt Oxia, 1750–1850 m, L.-Å.Gustavsson 6380 (LD); Macedonia, Pindos Mtns, Perivoli, Mt Aphtia, 

1700–2100 m, K.H.Rechinger 18516 (W). TURKEY-in-EUROPE: Istanbul, Silivri, Sinekli, Delice & Bulut 

(LD, ISTF 30535). 

Characterized by small, few-flowered, usually hemispherical clusters, papillose peduncles 

and relatively short seed appendages. Plants from E Bulgaria and the Sofia vicinity are 

±variable in the diagnostic characters, perhaps due to hybridization. In the S Pirin Mtns, 

introgression with L. campestris subsp. campestris was observed (J.Kirschner, Preslia 63: 

81–112, 1991). The distribution is imperfectly known. 

101

102 


54. Luzula taurica (V.I.Krecz.) Novikov, Nov. Sist. Vyssh. Rast. 27: 20 (1990) 

Luzula campestris subsp. taurica V.I.Krecz., Žurn. Russk. Bot. Obshch. Akad. Nauk S.S.S.R. 12: 490 (1928); 

Luzula multiflora subsp. taurica (V.I.Krecz.) Novikov, in Fl. Evr. Casti SSSR 2: 83 (1976). T: Crimea, 

Demerdzhi Yaila, 25 May 1905, N.A.Busch; lecto: LE, fide J.Kirschner, Taxon 39: 113 (1990). 

Luzula multiflora subsp. caucasica V.I.Krecz., Žurn. Russk. Bot. Obshch. Akad. Nauk S.S.S.R. 12: 490 (1928). 

T: [Georgia, Ahalcihe] ad fines inter Transcaucasiam rossicam et Turciae districtum Achalzich, 7 Jul 1874, 

G.F.Radde; syn: LE; [Georgia] Cartalinia, prope Turam [?], A.J.Szovits 25; syn: LE. 

Luzula campestris var. debilis Velen., Fl. Bulg. 572 (1891). T: Bulgaria, Mt Kom above Berkovica, Jul 

1889, J.Velenovský, syn: PRC, W. 

Luzula campestris var. atrofusca Maly, Glasn. Zemaljsk. Muz. Bosni Hercegovini 32: 141 (1920). T: Bosnia, 

Sarayevo, Mt Gola Jahorina, K.Maly; syn: SARA. 

Perennials, usually 15–25 cm tall, densely or rarely loosely caespitose; rhizome short, 

without stolons. Basal leaves usually up to 8–10 cm long, (2.5–) 3.0–4.0 (–5.0) mm wide; 

cauline leaves 1–2, usually 2.5–4.0 cm long, 2–3 mm wide; all leaves with obtuse tip and 

densely papillose-serrulate margins. Inflorescence composed of 1 (–3) subsessile and 2–6 (–7) 

pedunculate ovoid to hemispherical (5–) 7–9 (–11)-flowered clusters; peduncles straight, 

rarely slightly flexuose, ±smooth, without secondary branches; longest peduncles up to 2–4 cm 

long. Lower bract usually 1–3 cm long, shorter than inflorescence. Bracteoles membranous, 

1.5–2.5 mm long, ovate, fimbriate-ciliate above. Tepals ±equal, entire, acuminate, usually 

2.9–3.5 mm long, dark to blackish brown, with indistinct paler brownish margins. Stamens 6; 

anthers (0.8–) 1.1–1.3 (–1.4) mm long; filaments c. 0.5–0.6 mm long; style 0.7–1.1 mm long; 

stigmas 1.8–3.0 mm long, subpersistent. Capsule ±obovoid, ±shorter than perianth, pale 

brown to dark brown; capsule segments 2.4–3.0 mm long. Seeds 1.0–1.2 mm long, 0.7–0.8 mm 

wide; appendage (0.2–) 0.3–0.4 (–0.5) mm long. 2n=12 [12AL], fide J.Kirschner, Thaiszia 2: 

30–31 (1992). Fig. 22. 

Balkan Peninsula and Crimea in S Europe, Turkey, and Georgia and Armenia in the 

Caucasus. 13: ALB, BUL, GRC, ROM, TUE, YUC. 14: KRY. 33: NCS, TCS. 34: TUR. 

Subalpine meadows and grasslands. Map 85. 

13: ALBANIA, Skutari, Zukali, 15 Jul 1897, A.Baldacci [It. Alban. 5] 47 (K); Vermoshe,‘Gipfelregion von 

Grebeni Selces’, 1700 m, I.Dörfler 283 (G, K, M, WU). BULGARIA: Rila Mtns, Rilski Monastyr, Dolgia 

Rid, J.Štěpánková s.n. (PRA). GREECE: Serrai, Mt Vrondous, 1750–1849 m, S.Snogerup & A.Strid 5004 

(LD, PRA). ROMANIA: Hunedoara, Retezat, Mt Borescu, 12 Jul 1924, E.J.Nyárády et al. [Fl. Rom. Exs.] 

933 (K). F.Y.R. MACEDONIA: Vodno, 2 May 1972, R.Bělohlávková (PRA); Shar Planina, Kobilica, 2520 m, 

27 Aug 1938, B.Pawlowski (KRAM). 14: CRIMEA, Yalta, Nikitsky Yaila, 1350 m, P.H.Davis 33387 (K); 

Romak-Kosh, Babugan-Yaila, 20 Jul 1948, S.V.Yuzeptshuk (LE); Alushta, Luchistoe, Mt Demerdzhi, 

1100–1200 m, J.Kirschner L189 (PRA). 34: TURKEY: Trabzon, Karakö, N side of Pontus Pass, 2100 m, 

E.Hennipman et al. 1718 (K). 

Similar to L. abchasica and L. campestris. From the latter it differs in caespitose growth and 

seed characters. A very variable species (fide J.Kirschner, 1993: 170–172); the isolated 

populations differ from one another in a number of minor quantitative traits. 

55. Luzula divulgata Kirschner, Folia Geobot. Phytotax. 14 [1979]: 431 (1980) 

T: Czech Republic, Moravia, between Sobůlky and Věteřov, 6 May 1978, J.Kirschner s.n.; holo: PRC539. 

Illustration: J.Kirschner, Folia Geobot. Phytotax. 28: 159, fig. 6 (1993). 

Perennials, (10–) 15–25 (35) cm tall, densely caespitose, hairy; stolons absent; rhizome 

short. Basal leaves usually 10–15 cm long, 3.0–5.5 mm wide; cauline leaves usually 2 or 3, 

usually 3.5–6.5 cm long, 3–5 mm wide; all leaves with obtuse tip and sparsely papilloseserrate 

margin. Lower bract usually 1.6–3.0 cm long, shorter than inflorescence. 

Inflorescence composed of one subsessile and 2–8 pedunculate clusters; clusters broadly ovoid 

to hemispherical, 6–11-flowered. Peduncles straight or flexuose, smooth, without secondary 

branches. Longest peduncles (including clusters) 2.1–5.0 cm long. Bracteoles 1.8–2.4 mm 

long, fimbriate-ciliate above. Tepals ±equal, entire, ±acuminate, usually 3.5–4.7 mm long, 

castaneous brown to dark brown. Stamens 6; anthers usually 2.3–3.3 mm long; filaments 

0.5–0.7 mm long; ovary 1.0–1.7 mm long; style usually 1.7–1.9 mm long; stigmas 

subpersistent, 2.5–4.5 mm long, conspicuously exserted. Capsule brown, oblong-ovoid, not


Figure 22. Luzula multiflora subsp. multiflora. A mountain form. A, habit; B, flower cluster. 

Luzula campestris subsp. campestris. C, habit; D, seeds; E, perianth. Luzula taurica. 

F, habit. (A, B, Z.Skála & H.Skálová L338, PRA; C–E, J.Kirscher L501, PRA; 

F, J.Kirschner L189, PRA). Scale bars: A, C, F = 5 cm; B = 5 mm; D = 1 mm; E = 2 mm. 

Drawn by © E.Smrčinová. 

103

104 


mucronate, shorter than perianth; capsule segments (2.6–) 2.8–3.0 (–3.3) mm long. Seeds 

subglobular, 1.0–1.2 mm long, 0.9–1.0 mm wide; appendage 0.5–0.8 mm long. 2n=24 

[24AL], fide J.Kirschner, Folia Geobot. Phytotax. 14: 434 (1980); J.Kirschner, Thaiszia 2: 

20–22 (1992); J.Kirschner, Ann. Bot. Fenn. 29: 238 (1992). 

Europe, from E Sweden and N Poland, and Central Europe, to Bulgaria and Bosnia. 

10: SWE. 11: AUT, CZE, GER, HUN, POL. 13: BUL, ROM, YUC. 14: UKR. Open dry 

woodlands, clearings, forest paths. Map 86. 

10: SWEDEN: Uppland, Väddö, Edeby, H.Smith 3213 (K, KRA, LD, MA, W). 11: AUSTRIA: Wien, 

Neuwaldegg, 11 May 1905, J.Vetter (W). CZECH REPUBLIC: Bohemia, Rakovník, Nový Mlýn, Brabečka 

forest, J.Hašková, J.Kirschner & F.Krahulec JK222/89 (PRA). SLOVAKIA: Šahy, Dolné Turovce, 180 m, 

23 May 1959, J.Chrtek (LD). GERMANY: Bavaria, Regensburg, Irlbach, 20 Apr 1913, Schelling (M, PR). 

HUNGARY: Budapest, Mt Fazekashegy, 11 May 1919, A.Boros (BP). POLAND: Wolin Is., Międzyzdroje, 

L.Kirschnerová & J.Kirschner L324 (PRA); Krakow, Tyniec, 8 May 1923, K.Piech (KRAM). 

13: BULGARIA: Sofia, Gorni Lozen, Lozenska Planina Mtns, c. 1000 m, J.Kirschner L507, L508 (PRA). 

14. UKRAINE: Kharkov, Krasnokutsk, N.Tzvelev 8 (LE). 

Similar to L. multiflora subsp. multiflora in its ±robust caespitose growth but seeds 

±globose; appendage much longer and floral characters much closer to L. campestris. 

56. Luzula campestris (L.) DC., in J.B.A.P. de M. de Lamarck & A.P. de Candolle Fl. 

Franç., 3rd edn, 3: 161 (1805) 

Juncus campestris L., Sp. Pl. 329 (1753); Luzula campestris var. minor Bluff & Fingerh., Comp. Fl. German. 

1: 444 (1825); Luzula campestris α [unranked] vulgaris Gaudin, Fl. Helv. 2: 572 (1828), nom. inval.; Luzula 

campestris a. [unranked] genuina Asch., Fl. Brandenburg 740 (1864), nom. inval.; Gymnodes campestris (L.) 

Fourr., Ann. Soc. Linn. Lyon, sér. 2, 17: 172 (1869); Luzula vulgaris Buchenau, Bot. Jahrb. Syst. 7: 175 

(1885), nom. illeg.; Juncoides campestris (L.) Kuntze, Revis. Gen. Pl. 2: 724 (1891); Juncoides campestris 

var. vulgaris E.Sheld., Bull. Geol. Nat. Hist. Surv. 9: 64 (1894), nom. inval.; Luzula campestris subsp. 

vulgaris P.Fourn., Quatre Fl. France 153 (1935), nom. inval.; Luzula subpilosa [Gilib. ex] V.I.Krecz. & 

Gontsch., in V.L.Komarov, Fl. SSSR 3: 571 (1935), nom. illeg.; Luzula campestris f. vulgaris I.Grint., Fl. Rep. 

Soc. Roman. 11: 603 (1966), nom. inval. T: citation: ‘in pascuis Europae siccioribus’; lecto: LINN449.44, 

fide Carter, in Fl. Trop. E. Afr., Juncaceae, 8 (1966). 


Perennials with creeping rhizome and short stolons. Cauline leaves 2–3; all leaves with obtuse 

tip and margins papillose-serrulate. Inflorescence of 2–6 ovoid to hemispherical clusters; 

peduncles smooth, without secondary branches; longest peduncles usually 1.5–3.5 cm long 

(including clusters). Bracteoles 2–3 mm long, fimbriate-ciliate above. Tepals brown, 

castaneous brown or dark brown, equal, entire, acuminate, 2.8–4.2 mm long. Stamens 6; 

anthers 1.3–1.8 mm long; filaments 0.3–0.6 mm long; stigma subpersistent, 1.8–3.0 mm 

long. Capsules obovoid, equalling or ±shorter than perianth; capsule segments 2.1–3.2 mm 

long. Seeds 1.0–1.2 mm long, 0.8–1.0 mm wide; appendage 0.4–0.7 mm long. 

In most of its geographical range, L. campestris varies within very narrow limits. In the 

Iberian Peninsula, two subspecies are recognized. 

At least some peduncles nodding or flexuose 56a. subsp. campestris 

Peduncles straight, rigid 56b. subsp. nevadensis 

56a. Luzula campestris (L.) DC. subsp. campestris 

Luzula campestris var. elata Schltdl., Fl. Berol. 1: 203 (1823). T: [Germany, Berlin area], 1821, 

D.F.L.Schlechtendal; lecto: HAL69139, designated here. 

Luzula campestris var. pumila Schltdl., Fl. Berol. 203 (1823). T: [Germany, Berlin area], 1821, 

D.F.L.Schlechtendal [ut var. nana]; lecto: HAL69140, designated here. 

Luzula campestris var. collina G.Mey., Fl. Hanov. Excurs. 582 (1849); Luzula campestris f. collina (G.Mey.) 

Buchenau, in A.Kneucker, Cyper. Junc. Exs., no 60, c. 1899; Allg. Bot. Z. Syst. 6: 228 (1900). 

T: [Germany, Hannover, otherwise not designated; the common form] 

Luzula campestris subsp. longistyla Čelak., Prodr. Fl. Böhmen 1: 85 (1867). T: not designated.


Luzula campestris A.II.a.2. [unranked] luteola Asch. & Graebn., Syn. Mitteleur. Fl. 2: 522 (1904); Luzula 

campestris var. luteola (Asch. & Graebner) V.I.Krecz., Žurn. Russk. Bot. Obshch. Akad. Nauk S.S.S.R. 12: 490 

(1928). T: not designated. 

Luzula campestris var. insularis Briq., Prodr. Fl. Corse 1: 246 (1910); Luzula campestris proles insularis 

(Briq.) Rouy, Fl. France 13: 265 (1912); Luzula insularis (Briq.) Prain, Ind. Kew., suppl. 5: 157 (1921). 

T: Corse, entre Ste. Lucie et La Trinité, 50 m, 7 May 1907, J.I.Briquet; lecto: G, designated here. 

Luzula chabertii Rouy, Fl. France 13: 266 (1912); Luzula campestroides H.Lév., Bull. Acad. Int. Géogr. Bot., 

ser. 4, 25: 48 (1917), nom. illeg. T: [France] Forêt d’Fontainebleau dans les Ventes Bourbon, 16 May 1871, 

A.C.Chabert; lecto: FI, fide J.Kirschner, Taxon 39: 112 (1990). 

Luzula campestris var. rectirama Murr, Neue Übers. Bl-Pfl. Vorarlberg 46 (1923). T: [Austria] Nofels, 

Gisingen und Letzebühel b. Feldkirch, J.Murr; syn: n.v. 

Luzula campestris var. terana P.Monts., Anales Inst. Bot. Cavanilles 21(2): 492 (1964). T: Zamora, 

Sanabria, Tera R., Furnio de Lacillos, 1600 m, Jun 1945, T.M.Losa; holo: BCF. 

Luzula campestris subsp. iberica P.Monts., Anales Inst. Bot. Cavanilles 21(2): 492 (1964). T: Madrid, 

Sierra de Guadarrama, Peñalara, 25 May 1912, C.Vicioso; holo: BC63308. 

Illustrations: H.E.Hess, E.Landolt & R.Hirzel, Fl. Schweiz 1: 521 (1967); J.Kirschner, Folia Geobot. Phytotax. 

28: 153, fig. 3 (1993). 


Perennial, 5–25 cm tall, ±densely ciliate, with creeping rhizome and short stolons. Stem with 

numerous basal and 2–3 cauline ±flat leaves; upper leaves 2–4 cm × 1.7–3.3 mm, with 

sparsely papillose-serrulate margins. Clusters 3–9-flowered; peduncles erect, often ±flexuose, 

usually at least one nodding. Lower bract 1–3 cm long, shorter than inflorescence. Bracteoles 

2.0–2.5 mm long, whitish-membranous, ciliate-fimbriate. Tepals 2.8–4.1 mm long, 

castaneous brown to dark brown. Anthers 1.3–1.8 mm long, 3.0–4.5 times longer than the 

0.3–0.6 mm long filaments; style 1.0–1.5 mm long; stigma 1.8–3.0 mm long, ±persistent. 

Capsule ±equalling tepals; capsule segments 2.1–3.0 mm long, dark brown. Seeds 

subglobular, 1.0–1.1 × 0.8–1.0 mm; basal appendage 0.4–0.7 mm long. Mean length of 

middle stomata in upper cauline leaves 30–35 µm. 2n=12 [12AL], fide H.Nordenskiöld, 

Hereditas 37: 336–337 (1951); J.Kirschner, Pl. Syst. Evol. 200: 5 (1996). Fig. 22. 

Europe, except for the eastern part, and North Africa. Introduced to North America, New 

Zealand, Australia, Macaronesia, Falkland Is. 10: DEN, FIN, FOR, GRB, IRE, NOR, SWE. 

11: All. 12: BAL, COR, FRA, POR, SAR, SPA. 13: ALB, BUL, GRC, ITA, ROM, TUE, 

YUC. 14: BLR, BLT, RUC, RUN, RUW, UKR. 20: ALG, MOR. 21: azo, mdr. 50: nsw, tas, 

vic. 51: nzn, nzs. 72: nfl. 75: mas. 90: fal. Map. 87 [secondary occurrence not shown] 

11. GERMANY: Berlin, Jungfernhaide, 120 m, Lackowitz [A.Kneucker, Cyp. Junc. Exs.] 60 (GZU, PR). 

POLAND: Chrzanów, Płaza, A.Jasiewicz & Pałkowa [Fl. Polon. Exs.] 390 (WRSL). SWITZERLAND: 

Chur, Campodels, C.Coaz [J.Braun-Blanquet, Fl. Raet. Exs.] 611 (BC, G, GZU). 12. FRANCE: Seine-et- 

Oise, Forêt de Sénart, De Baire [Duffour, Soc. Franc.] 2242 (BC, P). PORTUGAL: Algarve, Sierra 

Monchique, 600 m, P.Montserrat et al. (JACA590668). SPAIN: Madrid, El Escorial de Guadarrama, 

15 May 1966, M.Mayor (MAF87300); Burgos, Lunada, Portillo de Lunada, 1500–1600 m, P.Montserrat 

(JACA173783). 14. BELORUSSIYA: Minsk, 12 May 1986, Tzvelev (LE). RUSSIA NORTHWEST: 

Pskov, V.Andreev 1683 (LE). 90. FALKLAND IS.: East Falklands, Port Stanley, Beatrice Cove, 10 Nov 

1945, Hamilton 12 (BM). 

Numerous literature records under the name L. campestris from outside Europe mostly refer 

to other species in the sect. Luzula. In Spain and Portugal, subsp. campestris is rather 

variable, particularly in the flower size, inflorescence appearance and length of stolons; in 

other regions, the variation is very limited. 

56b. Luzula campestris subsp. nevadensis P.Monts., Anales Inst. Bot. Cavanilles 

21(2): 492 (1964) 

T: Spain, Sierra Nevada, Puerto Lobo, 2100 m, 14 Jul 1923, P.Font Quer; holo: BC150865. 

Map: P.Montserrat, Anales Inst. Bot. Cavanilles 21(2): map 8 (1964). 

Perennial, usually 20–32 cm tall; rhizome long, creeping; stolons present. Basal leaves 

sparsely ciliate, c. 3.0–4.5 mm wide; cauline leaves 2–3, with upper ones 3.5–5.0 cm long, 

2.0–2.8 mm wide. Lower bract usually 1.2–2.0 cm long, shorter than inflorescence. Clusters 

105

106 


5–8-flowered; peduncles straight. Bracteoles 2.4–3.0 mm long. Tepals 3.5–4.2 mm long, 

brown to dark brown. Anthers 1.4–1.8 mm long; filaments 0.4–0.5 mm long; ovary 1.2 mm 

long; style 1.4–2.2 mm long; stigmas 2–3 mm long, subpersistent. Capsule segments 2.8–3.2 mm 

long. Seeds oblong-ovoid, 1.1–1.2 mm long, c. 0.9 mm wide; appendage 0.5–0.6 mm long. 

Mean length of middle stomata in upper cauline leaves 44–45 µm. Polyploid, fide J.Kirschner, 

Pl. Syst. Evol. 200: 5 (1996). 

Endemic to Spain. 12: SPA. Mountain grasslands and open woodlands. Map 88. 

12. SPAIN: Barcelona, Vallgorguina, ‘ad Torrentem Can Gras’, 250 m, P.Monserrat (JACA2347). 

An imperfectly known taxon, presumed to be a polyploid derivative of subsp. campestris. In 

addition to the holotype gathering, only a single additional specimen is known. However, its 

location, rather far from Sierra Nevada and in a different ecological situation, raises some 

doubts about its identity. 

57. Luzula sudetica (Willd.) Schult., Oesterr. Fl., 2nd edn, 1: 573 (1814) 

Juncus sudeticus Willd., Sp. Pl. 2: 221 (1799); Luzula nigricans Desv., J. Bot. (Desvaux) 1: 158 (1809), nom. 

illeg.; Luzula campestris γ [unranked] nigricans [Desv.] Gaudin, Fl. Helv. 2: 572 (1828); Luzula multiflora 

var. nigricans (Gaudin) W.D.J.Koch, Syn. Fl. Germ. Helv. 734 (1837); Luzula campestris c. [Spielart] 

alpestris G.Mey., Chloris Han. 572 (1836); Luzula campestris d. [Spielart] sudetica G.Mey., Fl. Hanov. 

Excurs. 582 (1849); Luzula multiflora var. sudetica (Willd.) Parl., Fl. Ital. 2: 307 (1852), nom. illeg.; Luzula 

campestris subsp. alpestris (G.Mey.) Čelak., Prodr. Fl. Böhmen 1: 85 (1867); Gymnodes sudetica (Willd.) 

Fourr., Ann. Soc. Linn. Lyon, sér. 2, 17: 173 (1869); Luzula campestris subsp. sudetica (Willd.) Čelak., Prodr. 

Fl. Böhmen 3: 749 (1881), nom. illeg.; Luzula multiflora subsp. sudetica (Willd.) Arcang., Comp. Fl. Ital. 713 

(1882), nom. illeg.; Juncoides campestris var. sudetica (Willd.) Coville, Contr. U. S. Natl. Herb. 4: 208 

(1893), nom. illeg.; Luzula erecta subsp. nigricans (Gaudin) Berher, in L.Louis, Fl. Vosges, 2nd edn, 216 

(1887), nom. illeg.; Luzula multiflora subsp. nigricans (Gaudin) Neuman, Sveriges Fl. 665 (1901), nom. illeg.; 

Luzula erecta proles nigricans (Gaudin) Rouy, Fl. France 13: 266 (1912). T: [Czech Republic/Poland], ‘in 

Sudetis’ [theKrkonoše Mts.]; lecto: B-WILLD 6837, fide J.Kirschner, Taxon 39: 113 (1990). 

Luzula campestris var. alpina Gaudin, Agrost. Helv. 2: 247 (1811) [doubtfully valid, probably validated later]. 

T: [Swiss Alps, otherwise not designated]. 

Luzula althii Herbich, Select. Pl. Galic. 12 (1836); Luzula campestris f. althii (Herbich) I.Grint., Fl. Rep. Soc. 

Roman. 11: 603 (1966). T: Ukraine, ‘prope Putila, in alpe Ploska prope Seletin, in Luczina’, F.Herbich; 

lecto: CZERN21173. [in herb. W authentic material with Juncus triglumis and another sheet with Luzula 

multiflora and the same label]. 

Illustrations: T.Szynal & J.Mądalski, Atlas Fl. Polsk. 2(2): 148 (1931); J.Kirschner & Lippert, Ber. Bayer. 

Bot. Ges. 65: Plate 4 (1995); J.Kirschner, Folia Geobot. Phytotax. 28: 169, fig. 9 (1993). 


Perennials, 10–25 (–35) cm tall, loosely caespitose; rhizome ascending to oblique, branched; 

stolons absent. Basal leaves sparsely ciliate, 1.5–4.0 mm wide; cauline leaves 2–3; upper leaves 

4.5–6.0 cm long, 1.4–3.0 mm wide; all leaves with densely papillose-serrate margin and apex 

obtuse. Lower bract usually 1.5–3.5 cm long, overtopping inflorescence. Inflorescence 

usually partly congested, of 3–10 ellipsoidal to ovoid clusters; clusters (5–) 7–11 (–15)flowered, 

with some or most clusters pedunculate; peduncles straight, smooth, usually 

without secondary branches, up to 1.0–3.5 cm long. Bracteoles 1.2–1.5 mm long, ciliate. 

Tepals unequal to ±equal, ±lanceolate, blackish brown, narrowly paler bordered above; outer 

tepals (1.9–) 2.1–2.5 (–2.7) mm long. Stamens 6; anthers (0.4–) 0.5–0.7 (–0.8) mm long; 

filaments 0.4–0.5 mm long; ovary 0.8–1.0 mm long; style 0.1–0.3 mm long; stigma early 

deciduous, 0.8–1.4 mm long. Capsules narrowly obovoid, subacute, indistinctly mucronate, 

equalling or shorter than perianth; capsule segments 1.7–2.0 × c. 1.0 mm. Seeds narrowly 

ellipsoidal, 0.9–1.0 mm long, 0.5–0.6 mm wide; appendage c. 0.1 mm long, fibrillate. 2n=48 

[48CL], fide J.Kirschner, Thaiszia, J. Bot. 2: 29–30 (1992); J.Kirschner & Lippert, Ber. 

Bayer. Bot. Ges. 65: 18 (1995); J.Kirschner, Pl. Syst. Evol. 200: 6 (1996). Fig. 23. 

Europe, from Iceland, Scandinavia and C European mountains to the Pyrenees, N Italy and 

the Balkans. 10: FIN, ICE, NOR, SWE. 11: AUT, CZE, GER, POL, SWI. 12: COR, FRA, 

SPA. 13: ALB, BUL, GRC, ITA, ROM, YUC. 14: RUN, UKR. Recorded from the Caucasus 

and Turkey but further study is needed. Montane and alpine moist meadows on peaty soils. 

Map 89.


Figure 23. Luzula sudetica. A, habit; B, flower cluster; C, D, outer and inner tepals; 

E, bracteole; F, flower; G, capsule; H, seed; I, stem T.S.; J, leaf T.S. (A-J, B.Błocki, 31 Jul 

1905, KRA). Scale bars: A = 5 cm; B = 5 mm; C, D = 1 mm; E = 1 mm; F, G = 2 mm; 

H = 1 mm; I, J = 1 mm. Drawn by T.Szynal. Reproduced with permission, from T.Szynal & 

J.Mądalski, Atlas Fl. Pol. 2/2: 148 (1931). 

107

108 


10. FINLAND: Savonia borealis, Lapinmäki, H.Lindberg [Pl. Finl. Exs.] 157 (H, LD, S, UPS, WRSL). 

11. CZECH REPUBLIC: the Krkonoše Mtns, Harrachov, Vosecká bouda, 1100–1130 m, 15 Jul 1998, 

L.Kirschnerová & J.Kirschner 4468 (PRA). GERMANY: Bayerischer Wald, summit of Mt Gr. Arber, 

28 Jul 1909, F.Vollmann (M). SLOVAKIA: Belianske Tatry, Kežmarská chata, J.Kirschner L301 (PRA). 

12. FRANCE: Ht. Pyrénées, Passades d’Aumur, Neouvielle, 2180 m, P.Montserrat (JACA275480). SPAIN: 

Huesca, Tramacastilla de Tena, El Bujicar, 1600 m, 27 Jul 1982, L.Villar (JACA129282). 

13. BULGARIA: Rila Mtns, Samokov, between ‘Komplex Maljovica’ and Maljovica chalet, 1900 m, 8 Aug 

1990, J.Štěpánek [K 409] (PRA). GREECE: Nom. Serrai, Mt Vrondous, 1400–1450 m, S.Snogerup & 

A.Strid 5015 (LD). ITALY: Piemont, Valli di Lanzo, Lac Falin, 1600 m, 21 Jul 1987, A.Pistarino (BM). 

ROMANIA: Retezat Mtns, Lake Zanoga, 7 Aug 1896, Wagner (LD). 14. UKRAINE: the Carpathians, 

Vorokhta, 1430 m, V.M.Vinogradova & Stankova [Gerb. Fl. SSSR] 5910 (LE). 

Towards the south, most plants have inner and outer tepals ±of the same length, otherwise 

the variation limits are narrow. The loosely caespitose growth with oblique/ascending 

branched rhizome is an important diagnostic character. 

58. Luzula congesta (Thuill.) Lej., Fl. Spa 1: 169 (1811) 

Juncus congestus Thuill., Fl. Env. Paris, 2nd edn, 178 (1800); Luzula erecta var. congesta (Thuill.) Desv., 

J. Bot. (Desvaux) 1: 156 (1809); Luzula campestris var. congesta (Thuill.) Bicheno, Trans. Linn. Soc. London 

12: 334 (1819); E.Mey., Syn. Luzul. 18 (1823); Luzula intermedia var. congesta (Thuill.) Spenn., Fl. Friburg. 

1: 178 (1825); Luzula multiflora var. congesta (Thuill.) W.D.J.Koch, Syn. Fl. Germ. Helv. 734 (1837); Luzula 

campestris subsp. congesta (Thuill.) K.Richt., Pl. Eur. 1: 186 (1890); Luzula multiflora subsp. congesta 

(Thuill.) Arcang., Comp. Fl. Ital. 713 (1882); Luzula multiflora f. congesta (Thuill.) Neuman, Sveriges Fl. 665 

(1901); Luzula erecta proles congesta (Thuill.) Rouy, Fl. France 13: 265 (1912). T: [France, Paris], 

J.L.Thuillier; lecto: G-DEL, fide J.Kirschner, Taxon 39: 110 (1990); isolecto [or other authentic material]: K. 

Juncus campestris var. strictus Retz., Fl. Scand. Prodr., 2nd edn, 82 (1795). T: 'Gramen hirsutum capitulo 

globoso. Mor. 3. s. 8. t. 9'; R.Morison, Pl. Hist. Univ. Oxon. 3: 224-225, sect. 8, tab. 9 [unnumbered figure, 

first in upper left hand corner] (1715); holo. not located [no material in herb. Retzius, LD]. 

Juncus liniger With., Syst. Arr. Brit. Pl., 4th edn, 2: 343 (1801). T: England,‘Shortwood near Pucklechurch, 

Swayne; holo: not extant. 

Luzula spicata var. latifolia Losa, Contrib. Est. Fl. Alava, Vitoria 1946: 43 (1946), n.v. T: Spain, Sierra de 

Cantabria, Alava, Lagrán, Bujumendia, Jul 1928, T.M.Losa; syn: BCF, Jun 1933, T.M.Losa; syn: BCF, n.v. 

Luzula campestris var. pallens Merino, Mem. Real Soc. Esp. Hist. Nat. 2: 472 (1904), nom. illeg., non Asch. & 

Graebn. (1904). T: Spain, Galicia, Galdo, Rodríguez Franco; holo, n.v., fide P.Montserrat, Anales Inst. Bot. 

Cavanilles 21: 507 (1964). 

Illustrations: F.G.P.Buchenau, in H.G.A.Engler, Pflanzenr. (iv.36) 25: 91, fig. 54 (1906); J.Kirschner, Folia 

Geobot. Phytotax. 28: 156,157, fig. 4, 5 (1993). 


Perennials, 25–55 cm tall, densely caespitose; rhizome ±short, vertical; stolons absent. Basal 

leaves densely ciliate, 3.5–6.0 mm wide; cauline leaves usually 3; upper leaves 2.5–6.5 cm 

long, 1.5–4.0 mm wide; all leaves with obtuse tip and margins papillose-serrulate. Lower 

bract usually 1.5–3.0 cm long, overtopping or less often shorter than inflorescence. 

Inflorescence usually a congested compact, ovoid to ±pyramidal head, of 4–8 clusters; 

clusters usually 7–10-flowered, sometimes some clusters pedunculate; peduncles straight, up 

to 0.9–3.0 cm long, ±smooth. Bracteoles 2.5–2.8 mm long, fimbriate-ciliate. Tepals ±equal, 

glabrous, ±lanceolate, acuminate, pale straw-brown or brown; margins paler to membranous 

above, usually 2.9–3.5 mm long. Stamens 6; anthers (0.6–) 0.7–0.9 mm long; filaments 

0.6–0.8 mm long; ovary 1.0–1.2 mm long; style 0.5–0.8 mm long; stigmas 2.0–2.4 mm long. 

Capsules obovoid to subglobular, pale brown, ±obtuse, equalling or shorter than perianth; 

capsule segments 2.3–3.0 × 1.5–1.8 mm. Seeds ovoid or oblong-ovoid, 1.2–1.5 mm long, 

(0.8–) 0.9–1.0 mm wide; appendage 0.4–0.6 mm long, usually fibrillate. 2n=48 [48AL], fide 

H.Nordenskiöld, Hereditas 42: 10–11 (1956); J.Kirschner, Thaiszia, J. Bot. 2: 19–20 (1992); 

J.Kirschner, Pl. Syst. Evol. 200: 8 (1996). 

Western Europe, from W and S Scandinavia and Germany to Spain and Portugal. Introduced 

to Madeira, Azores, New Zealand and Australia. 10: DEN, FOR, GRB, ICE, IRE, NOR, SWE. 

11: BGM, GER, NET, POL. 12: FRA, POR, SPA. 21: azo, mdr. 50: nsw, tas. 51: nzn, nzs. 

Peat-bogs, wet meadows on peaty soils, from sea level to c. 1800 m. Map 90.


10. DENMARK: Jylland, Varde, 29 Jul 1951, H.Nordenskiöld 51-74 (UPS). GREAT BRITAIN: Isle of 

Man, W of Ballaweanagh, S of Sarwick, 25 Jun 1950, V.S.Summerhayes (K). IRELAND: Co. Carlow, 

Bagnelstown, T. C. G. Rich 4 [L256] (PRA). NORWAY: Hordaland, Eldöyane, 16 Aug 1955, J.Ericson 

(LD). 11. BELGIUM: Flandre, Wingene, 8 Jun 1979, Stieperaere [Soc. Éch. Pl. Vascul. Eur. Occid.] 9906 

(LD, LG). GERMANY: Niedersachsen, Oldenburg, Jaderberg, 3 Jul 1920, F.Müller (LD). POLAND: 

Kołobrzeg, Dogowo, J.Mądalski 13729 (WRSL, herb. Mądalski). 12. SPAIN: Navarra, Selva de Irati, 

Ochagavia, 950 m, G.Montserrat et al. (JACA121187). PORTUGAL: Coimbra, Carregaes d’Eiras, May 

1896, M.Ferreira [Fl. Lusit. Exs.] 1417 (LD). 21. AZORES: São Miguel, Ribeira Grande, Lagoa do Fogo, 

14 May 2000, J.Kirschner (PRA). 

59. Luzula multiflora (Ehrh.) Lej., Fl. Spa 1: 169 (1811), nom. cons. prop. 

Juncus campestris var. multiflorus Ehrh., Beitr. Naturk. 5: 14 (1790); Juncus multiflorus (Ehrh.) Hoffm., 

Deutschl. Fl., Dritter Jahrg., 1: 169 (v–xii 1800), nom. illeg., non Retz. (1795), nec Desf. (1799); Luzula 

intermedia var. multiflora (Ehrh.) Spenn., Fl. Friburg. 1: 178 (1825); Luzula campestris subsp. multiflora 

(Ehrh.) Schübl. & M.Martens, Fl. Würtemberg 230 (1834); Gymnodes multiflora (Ehrh.) Fourr., Ann. Soc. 

Linn. Lyon, sér. 2, 17: 173 (1869); Juncoides campestris var. multiflora (Ehrh.) E.Sheld., Bull. Geol. Nat. Hist. 

Surv. 9: 64 (1894); Juncoides multiflora (Ehrh.) Druce, Ann. Scott. Nat. Hist. 1904: 42 (1904). T: Italy, 

S Tirol, Mendelgebirge, between St Pauls and Perdonig, 46°29'N 11°15'E, I.Hertel & H.Hertel; typus cons. 

prop.: M33725; iso: PRA (2n=36, karyotype 36AL). 

Luzula campestris subsp. carpetana Rivas Mart., Anales Inst. Bot. Cavanilles 21/1: 288 (1963). T: [Spain, 

Sierras de Guadarrama y Gredos], S.Rivas Martínez; holo: MAF, n.v. 

Luzula jilongensis K.F.Wu, J. E. China Norm. Univ., Nat. Sci. Ed., 1992(3): 95 (1992). T: Tibet, Jilong 

Xian, 3400–3800 m, 5 Jul 1975, Cheng-yih Wu & Shu-kun Chen 75-612 (KUN). 

Luzula multiflora var. intermedia Koidz., Pl. Sachal. Nak. 32, fig. 1 (1910), n.v.; Luzula multiflora var. tenuis 

Satake, in T.Nakai & M.Honda, Nov. Fl. Jap. 1: 36 (1938). T: [Russia] Sakhalin, Dobky, 24 Jun 1906, 

G.Nakahara; lecto: TI [as L. campestris var. intermedia], designated here. 

Map: E.Hultén, Kongl. Svenska Vetenskapsakad. Handl., ser. 5, 8(5): map 64 (1962). 

Perennials, ±densely caespitose; rhizome shor t; stolons absent. Basal leaves densely ciliate 

to subglabrous, 2.5–4.0 (–6.0) mm wide; cauline leaves 1–3 (–4), with densely papilloseserrulate 

margins. Lower bract leaf-like, usually longer than or equalling inflorescence (less 

often shorter). Inflorescence usually composed of pedunculate clusters, or less often clusters 

subcongested; peduncles smooth or papillose. Bracteoles scarious, to 2 mm long, subentire to 

ciliate-fimbriate. Tepals ±equal to subequal, acuminate to subaristate, 2.4–3.5 (–3.9) mm 

long, straw, straw-brown, castaneous-brown to dark brown, rarely blackish, usually with a 

distinct paler margin. Stamens 6; anthers usually 0.6–1.0 (–1.4) mm long, anther/filament 

ratio usually 1.0–1.5 (–2.5); style 0.3–0.7 (–0.8) mm long; stigmas early deciduous, 

1.1–2.0 (–3.1) mm long. Capsule obovoid, smooth, shorter to slightly longer than perianth, 

pale brownish to dark brown; capsule segments 1.9–2.8 mm long. Seeds oblong-ovoid to 

ellipsoid-ovoid, 0.8–1.2 mm long, 0.6–0.9 mm wide; appendage 0.2–0.4 (–0.5) mm long. 

An extremely widespread and variable polyploid, rarely (partially) agmatoploid, species. 

In some regions (China, USA, the Far East) it is not known satisfactorily, and additional taxa 

may be revealed. Six subspecies are recognized. 

1 Peduncles ±densely papillose throughout 59d. subsp. snogerupii 

1: Peduncles smooth, rarely with scattered papillae just below clusters 

2 Peduncles sometimes (usually one or more in a plant) recurved 59e. subsp. hibernica 

2: Peduncles straight to (rarely) flexuose, not recurved 

3 Seeds (excluding appendage) (0.8–) 0.9 mm long; tepals to 

2.7 mm long; capsule often longer than perianth; seed appendage 

(0.3–) 0.4 mm long 59f. subsp. monticola 

3: Seeds (0.9–) 1.0–1.1 (–1.2) mm long; tepals usually longer than 

2.7 mm, or if tepals to 2.7 mm long and capsule exceeding perianth, 

then seed appendage 0.2–0.3 mm long 

109

110 


4 Capsule subacuminate, usually distinctly exceeding perianth; style 

0.3–0.5 mm long 59b. subsp. frigida 

4: Capsule subobtuse, usually shorter than perianth; style 

(0.4–) 0.5–0.8 mm long 

5 Seed appendage c. 0.3 mm long; capsule segments 1.9–2.2 mm 

long 59c. subsp. sibirica 

5: Seed appendage usually 0.4 mm long; capsule segments 

(2.0–) 2.3–2.6 (–2.8) mm long 59a. subsp. multiflora 

59a. Luzula multiflora (Ehrh.) Lej. subsp. multiflora 

Juncus intermedius Thuill., Fl. Env. Paris, 2nd edn, 178 (1800); Juncus erectus Pers., Syn. Pl. 1: 386 (1805), 

nom. illeg.; Luzula erecta Desv., J. Bot. (Desvaux) 1: 156 (1808), nom. illeg.; Luzula intermedia (Thuill.) 

Spenn., Fl. Friburg. 1: 177 (1825), nom. illeg., non Nocca & Balb. (1816); Juncoides erecta Boulger, in 

C.A.Johns, Flowers of Field, 29th edn, 755 (1899); Juncoides intermedia (Thuill.) Rydb., Bull. Torrey Bot. 

Club 32: 610 (1905); Luzula campestris subsp. erecta [Desv.] P.Fourn., Quatre Fl. France 153 (1935). 

T: [France, Paris], J.L.Thuillier; lecto: G-DEL, fide J.Kirschner, Taxon 39: 110 (1990). 

Juncus campestris var. tenuis Retz., Fasc. Obs. Bot. 12 (1774). T: [without locality and collector] herb. 

Retzius; holo: LD. 

Juncus nemorosus Host, Icon. Descr. Gram. Austriac. 3: 64 (1805), nom. illeg., non Pollich (1776); Luzula 

nemorosa Hornem., Hort. Bot. Hafn. 962 (1815); Luzula campestris δ [unranked] nemorosa (Hornem.) 

Gaudin, Fl. Helv. 2: 572 (1828); Luzula campestris var. nemorosa (Hornem.) Neilr., Nachtr. Fl. Wien 101 

(1851). T: icon in N.T.Host, Icon. Descr. Gram. Austriac. tab. 97, fig. 5 (1805); lecto, fide J.Kirschner, 

Taxon 39: 111 (1990). 

Luzula spicata var. tenella [Miel. ex] E.Mey., Linnaea 22: 415 (1849); Luzula tenella ([Miel. ex] E.Mey.) 

R.Hinterh. & J.Hinterh., Prodr. Fl. Salzburg 223 (1851); Luzula multiflora subsp. tenella (E.Mey.) 

Kožuharov, Izv. Bot. Inst. (Sofia) 11: 135 (1963). T: [Austria] Schattbachalpen in Grossarl, M.Mielichhofer; 

lecto: REG85873, fide J.Kirschner, Folia Geobot. Phytotax. 28: 160 (1993). 

Luzula multiflora var. uliginosa Gremli, Excursionsfl. Schweiz, 3rd edn, 377 (1878). T: [Switzerland, 

otherwise not designated]. 

Luzula intermedia Figert, Deutsche Bot. Monatsschr. 15: 12 (1897), nom. illeg., non Thuill. (1800); Luzula 

ambigua Sóo, Acta Bot. Acad. Sci. Hung. 16: 367 (1971), pro hybr. T: [Poland, Legnica] Schlesien, 

Liegnitz [several specimens from the vicinity of the town cited, material not found]. 

Luzula multiflora var. flexuosa Beyer, Verh. Bot. Vereins Prov. Brandenburg. 41: xxiii (1900). 

T: [Switzerland] Graische Alpen, Beauregard, Val Grisanche, 18 Jul 1889, R.Beyer ; lecto: W, designated here. 

Luzula campestris [Rasse] pallens Asch. & Graebn., Syn. Mitteleur. Fl. 2(2): 524 (1904), an nom. illeg., non 

Merino (1904); Luzula campestris var. pallescens Husn., Joncées 26 (1908), nom. illeg.; Luzula multiflora f. 

umbrosa Neuman, Sveriges Fl. 665 (1901); Luzula erecta var. pallens (Asch. & Graebn.) Rouy, Fl. France 13: 

265 (1912). T: icon in D.H.Hoppe, Deutschl. Fl. 1/77: sine no. (1839); lecto: designated here. 

Luzula erecta proles pyrenaica Sennen, Bull. Soc. Bot. France 63: 135 (1916); Luzula multiflora subsp. 

pyrenaica (Sennen) P.Monts., Anales Inst. Bot. Cavanilles 21(2): 501 (1964) [uncertain validity]. 

T: France, Cerdagne au Cambredase, E.M.Sennen [Pl. Esp.] 2359; syn: BC, BM, W. 

Luzula campestris var. acadiensis Fernald, Rhodora 19: 38 (1917); Luzula multiflora var. acadiensis (Fernald) 

Fernald, Rhodora 47: 267 (1945). T: Canada, Prince Edward Is., Charlottetown, W side of St Peters Bay, 

30 Jun 1914, M.L.Fernald & H.St.John [Pl. Exs. Gray.] 190; syn: GH, S. 

Luzula multiflora subsp. occidentalis V.I.Krecz., Žurn. Russk. Bot. Obshch. Akad. Nauk S.S.S.R. 12: 490 

(1928). T: not designated (not located at LE). 

Illustrations: T.Szynal & J.Mądalski, Atlas Fl. Polsk. 2(2): 149 (1931); H.E.Hess, E.Landolt & R.Hirzel, 

Fl. Schweiz 1: 521 (1967); J.Kirschner, Folia Geobot. Phytotax. 28: 163, fig. 7 (1993). 

Map: E.Hultén, Kongl. Svenska Vetenskapsakad. Handl., ser. 5, 8(5): map 64 (1962). 

Stems 15–38 cm tall. Basal leaves densely to sparsely ciliate, 3–6 mm wide; cauline leaves 

usually 2–4, 3.7–12 cm long, 1.8–5 mm wide. Lower bract usually 2.5–6.5 cm long, 

exceeding inflorescence or shorter. Inflorescence usually of (4–) 5–12 (–17) pedunculate 

clusters; clusters ellipsoid to ovoid, usually (6–) 8–14 (–18)-flowered; peduncles to 1.5–5.2 

cm long, straight, ±smooth, rarely with secondary branches. Bracteoles 1.5–2.5 mm long, 

subentire. Tepals pale yellowish-greenish to dark brown, 2.5 to 3.9 mm long. Anthers 

0.7–1.4 mm long, anther/filament length ratio 1.0–2.0 (–2.5); style 0.4–0.8 mm long; stigma


1.3–3.1 mm long. Capsule obovoid to subglobular, equalling perianth or shorter; capsule 

segments (2.0–) 2.2–2.6 (–2.8) mm long. Seeds ovoid to ellipsoid, 0.9–1.2 mm long, 

(0.6–) 0.7–0.8 (–0.9) mm wide; appendage 0.3–0.4 (–0.5) mm long. 2n=24 [24AL], 36 

[36AL], fide J.Kirschner, Thaiszia 2: 23–27 (1992). Fig. 22, 24. 

An extremely variable subspecies occurring very frequently in innumerable forms in most of 

Europe, many regions of West Asia, in North America (partly introduced), and introduced in 

a number of regions (more frequently in Australia, New Zealand, rarely in many additional 

regions). Many literature records refer to other taxa (particularly from W North America, SE 

Asia etc.). Only regions where the occurrence was safely confirmed on the basis of good 

herbarium material are listed here. It is difficult to distinguish the secondary occurrence from 

native distribution. 10: DEN, FIN, FOR, GRB, ICE, IRE, NOR, SWE. 11: All. 12: COR, 

FRA, POR, SPA. 13: ALB, BUL, GRC, ITA, ROM, TUE, YUC. 14: All. 20: ALG, MOR. 21: 

azo, cny, mdr. 30: WSB. 31: KHA, SAK. 32: KAZ, KGZ, TKM, TZK, UZB. 33: NCS, TCS. 

34: IRN, TUR. 36: CHC, CHT. 38: JAP, KOR, tai. 40: WHM, EHM. 50: tas. 51: nzn, nzs. 

70: GNL. 71: ABT, BRC, MAN. 72: LAB, NBR, NFL, NSC, ONT, PEI, QUE. 73: MNT, 

ORE. 74: IOW, MIN, NDA, SDA. 75: CNT, INI, MAI, MAS, MIC, NWH, NWJ, NWY, 

OHI, PEN, VER, WVA. 78: DEL, VRG. 80: COS. 85: AGE. 90. FAL. In a variety 

of habitats, most often in meadows (usually wet) and montane to subalpine grasslands, open 

woodlands, usually disturbed, road verges etc. from sea level to subalpine altitudes. 

Map 91 (primary and secondary distributions not distinguished on the map). 

11. GERMANY: Westfalen, Hagen, Philipshöhe, 16 May 1871, P.E.E.Sintenis (LD). 12. FRANCE: Loir-et- 

Cher, La Ferté-Imbault, 22 May 1924, L.Segret [Duffour, Soc. Franc.] 4815 (P); Gorges de Bitet, Laruns, 

Ossau, 1400 m, 5 Aug 1980, P.Montserrat & L.Villar (JACA330180). SPAIN: Andorra, Val del Riu, 1948, 

T.M.Losa (BCF4093); León, Las Omanas, 12 May 1973, M.Pérez (MA488994). 14. RUSSIA CENTRAL: 

Moscow, Pushkino, Komyagino, 10 Jul 1974, G.M.Proskuryakova (W). RUSSIA NORTH: Komi, Ust- 

Kulomskyi Distr., B. Soplesa R., 1 Aug 1928, S.Naumova 344 (MW). 21. AZORES: Fayal, Falca, 13 Apr 

1962, M.L.Gonçalves 297 (BM). 72. QUÉBEC: Sherbrooke, 10 Jun 1976, S.Brisson 76234 (LD). 

30. WEST SIBERIA: Arctic Urals, 4 Aug 1926, V.B.Sochava 402 (LE). 

59b. Luzula multiflora subsp. frigida (Buchen.) V.I.Krecz., Žurn. Russk. Bot. Obshch. 

Akad. Nauk S.S.S.R. 12: 490 (1928) 

Luzula campestris var. frigida Buchenau, Oesterr. Bot. Z. 48: 184 (1898); Luzula frigida (Buchenau) Sam., in 

C.A.M.Lindman, Sv. Fanerogamfl. 161 (1918); Luzula sudetica var. frigida (Buchenau) Fernald, Rhodora 47: 265 

(1945); Luzula kjellmaniana subsp. frigida (Buchenau) Schljakov, Novosti Sist. Vyssh. Rast. 16: 85 (1979). 

T: Norway, Kongsvold, Dovre Fjeld, Drivathal, 3 Aug 1892, K.G.Baenitz [Herb. Eur.] 7120; lecto: PR, fide 

J.Kirschner, Taxon 39: 113 (1990); isolecto: W. 

Luzula multiflora var. contracta Böcher, Meddel. Grønland 147(7): 17 (1950), nom. inval. [interpreted 

according to authentic material in C] 

Illustrations: J.Lid, Norsk & Svensk Fl. 199, fig. 98f, 203, fig. 100g (1963). 

Maps: E.Hultén, Kongl. Svenska Vetenskapsakad. Handl., ser. 5, 8(5): map 64 (1962); E.Hultén & M.Fries, 

Atlas N. Eur. Vasc. Pl. 1: map 173 (1986). 

Stems 13–30 cm tall. Basal leaves sparsely ciliate or subglabrous, 2.0–3.5 mm wide; cauline 

leaves usually 1–3, with upper ones 2.0–4.5 cm long and 1.0–2.8 mm wide. Lower bract 

usually 1.0–2.5 cm long. Inflorescence congested, of 2–6 clusters; clusters ellipsoid–ovoid or 

ovoid, 6–10-flowered, mostly pedunculate. Peduncles straight, smooth, rarely with secondary 

branches, to 0.8–2.5 cm long. Bracteoles 1.2–1.6 mm long, ciliate. Tepals dark brown, 

2.4–2.9 mm long. Anthers 0.6–0.9 mm long, anther/filament length ratio 0.8–1.2; ovary 

0.8–0.9 mm long; style 0.3–0.5 mm long; stigma 1.2–1.3 mm long. Capsules subacuminate, 

equalling or overtopping the perianth; capsule segments 2.0–2.4 mm long. Seeds ±ellipsoid, 

1.0–1.1 mm long, 0.7 mm wide; appendage 0.2–0.3 mm long. 2n=36 [36AL], fide 

J.Kirschner, Thaiszia 2: 27 (1992). 

Northern Europe, Iceland, Greenland, distribution in North America imperfectly known, but 

certainly from Alaska and Canada, and probably also in the mountains of NE USA. Very 

similar plants from the Far East require further investigation. 10: FIN, ICE, NOR, SWE. 14: 

111

112 


Figure 24. Luzula multiflora subsp. multiflora. A common lowland form. A, habit; B, flower 

cluster; C, flower; D, capsule; E, tepal; F–H, seeds; I, stem T.S.; J, leaf T.S. 

(A–F, I, J, B.Błocki, 28 May 1898, KRA; G, J.Mądalski s.n., 1928, KRA). Scale bars: 

A = 5 cm; B = 5 mm; C–H = 2 mm; I, J = 1 mm. Drawn by T.Szynal. Reproduced with 

permission, from T.Szynal & J.Mądalski, Atlas Fl. Pol. 2/2: 149 (1931).


RUN, RUW. 30: WSB. 70: ASK, GNL, NWT, YUK. 71: ABT, BRC, MAN, SAS. 72: LAB, 

NFL, NSC, ONT, QUE. 75: MAI, MAS, NWH, VER. Subarctic and mountain tundra, peatbog 

margins, meadows. Map 92. 

10. FINLAND: Inari, Angeli Station, 13 Jul 1970, Kause & Seikula (M). NORWAY: Hedmark, Folldal, 

840–900 m, 19 Aug 1965, K.P.Buttler & Gauhl (M); Lyngenfjord, Lyngseidet, 26 Jul 1988, K.G.Baenitz (M). 

SWEDEN: Karesuando, Gunnarin Korva, S.O.Björkman 3010 (UPS, 2n=36); Lule Lappmark, Avvako, 550 m, 

21 Jul 1967, H.Hertel (M); Härjedalen, Tännäs, Funäsdalen, 21 Jul 1916, M.Östman (BRNU). 14. RUSSIA 

NORTH: Murmansk, Poyakonda, M.Šourková 6871 (PRA). 71. MANITOBA: Churchill, K.H.Rechinger 

1375 (W). 72. QUEBEC: Great Whale R., Poste-de-la-Baleine, 30 Jul 1969, S.Brisson & Forest (LD). 

Intermediates are known between subsp. frigida and subsp. multiflora. Plants apparently 

close to L. multiflora subsp. frigida are sometimes recorded in Asia (China, Mongolia); 

similar plants are occasionally found in C Europe [J.Kirschner, Preslia 67: 219–223 (1996)]; 

they are treated as a part of the variation range of subsp. sibirica(Chinese plants) and subsp. 

multiflora (C European plants). 

59c. Luzula multiflora subsp. sibirica V.I.Krecz., Žurn. Russk. Bot. Obshch. Akad. Nauk 

S.S.S.R. 12: 490 (1928) 

Luzula sibirica (V.I.Krecz.) V.I.Krecz., in B.A.Fedtschenko, Fl. Transbaic. 2: 144 (1931); Luzula 

kjellmaniana subsp. sibirica (V.I.Krecz.) Schljakov, Novosti Sist. Vyssh. Rast. 16: 85 (1979). T: Russia, 

Siberia, Kitoi, Osna R., Cherski & Hartung 494; lecto: LE, fide J.Kirschner, Taxon 39: 113 (1990). 

Luzula multiflora subsp. asiatica Krylov & Serg., in P.N.Krylov, Fl. Zap. Sibiri 3: 556 (1929). T: not 

designated. 

Luzula multiflora f. conglobata Krylov & Serg., in P.N.Krylov, Fl. Zap. Sibiri 3: 556 (1929). T: not 

designated. 

Luzula multiflora f. umbelliformis Krylov & Serg., in P.N.Krylov, Fl. Zap. Sibiri 3: 556 (1929). T: not 

designated. 

Luzula multiflora var. minuta Krylov & Serg., in P.N.Krylov, Fl. Zap. Sibiri 3: 556 (1929). T: numerous 

syntypes from the Altai and Semipalatinsk alpine regions, otherwise not indicated. 

Luzula vinogradovii Sipliv., Novosti Sist. Vyssh. Rast. 12: 290 (1975). T: Russia, Chita Region, Kyra, 

Sokhondo, 3 Jul 1972, A.B.Vinogradov & V.K.Siplivinski; holo: LE. 

Luzula multiflora var. kjellmannioides R.L.Taylor & MacBryde, Canad. J. Bot. 56(2): 191 (1978). 

T: Russia, S Kamchatka, 3 Jul 1921, E.Hultén 1924; holo: S. 

Luzula badia K.F.Wu, J. E. China Norm. Univ., Nat. Sci. Ed., 1992(3): 99 (1992). T: China, Xinjiang, 

7 Jul 1967, anonymus 6275; holo: XJBRI. 

Map: N.K.Kovtonyuk, in L.I.Malyshev & G.A.Peshkova, Fl. Sibir. 4 (Araceae – Orchidaceae): map 41 (1987) 

[Siberia]. 

Plants 12–35 cm tall. Basal leaves sparsely ciliate or subglabrous, 2.5–4.0 mm wide; cauline 

leaves usually 1 or 2; upper cauline leaves 3.5–7.0 cm long, 2.0–3.0 mm wide. Lower bract 

usually 2.5–4.5 cm long, overtopping inflorescence. Inflorescence subcongested with some 

pedunculate clusters or with most clusters pedunculate; clusters 3–7, ellipsoid-ovoid 

or ovoid, 7–10-flowered. Peduncles straight, smooth, rarely with secondary branches, to 

1.5–4.5 cm long. Bracteoles 1.5 mm long, subentire or ciliate. Tepals brown or dark brown, 

2.4–2.8 mm long. Anthers 0.6–0.9 mm long, anther/filament length ratio 1.0–1.3; ovary 

0.9 mm long; style 0.5–0.8 mm long; stigmas 1.2–1.5 mm long. Capsules obovoid to 

subglobular, equalling perianth or shorter; capsule segments 1.9–2.2 mm long. Seeds oblongovoid 

or ellipsoid, 1.0–1.1 mm long, 0.7 mm wide; appendages 0.3 mm long. 2n=36 [36AL] 

or 2n=24 [24AL], fide J.Kirschner, Thaiszia 2: 27(1992). 

Siberia and the Russian Far East, from the Urals to Kamchatka, parts of Middle Asia, 

N China and Mongolia. Probably also in Kazakhstan. 30: ALT, BRY, CTA, IRK, KRA, TVA, 

WSB, YAK. 31: AMU, KAM, KHA, KUR, MAG, PRM, SAK. 32: KAZ, KGZ. 36: CHI, 

CHN, CHX. 37: MON. Mountain grasslands, tundra, along streams and paths. Map 93. 

30. ALTAY: Usk-Koksu, Multa R., 1900 m, I.M.Krasnoborov 245 (NS); Yabogan, Yaboganskiy Pereval, 1850 

m, J.Kirschner 22 (PRA). CHITA: Borstshovochnyi Khrebet, Malaya Ashinga R., 12 Jul 1966, 

M.Maximova [Gerb. Fl. SSSR] 5053 (BRNU, LE). YAKUTIYA: Kolyma R. basin, Cherskii, 10 Aug 1983, 

S.Bubnova (NS, PRA); Neryungrinsk, Nagornyi, N.K.Kovtonyuk 1030 (NS, PRA). 31. MAGADAN: 

113

114 


Chukotka, E coast of Chaynskaya Guba, V.Petrovskiy & P.Zhukova 434 (LE, 2n=24); Chukotka, Bezimyannyi, 

3 Aug 1974, V.Petrovskyi et al. (LE, 2n=36). KHABAROVSK: Sofiisk, Bureika R., 1500 m, 10 Jul 1978, 

A.Nechaev (NS, PRA). 

Populations from Kamchatka and the Kuriles have an intermediate appearance between 

subsp. frigida and subsp. sibirica. In addition, plants close to the type subspecies are found 

in the southern part of the range of subsp. sibirica. 

59d. Luzula multiflora subsp. snogerupii Kirschner, Ann. Bot. Fenn. 29: 239 (1992) 

T: Greece, Nom. Serrai, Mt Vrondous, 2 km SW of the summit, 1250–1300 m, 14 Jul 1987, S.Snogerup, 

cultivated as L132; holo: PR; iso: LD, PRA. 

Plants 10–32 cm tall. Basal leaves sparsely ciliate, 2.5–4.0 mm wide; cauline leaves usually 2 

or 3, with upper ones 3.0–6.5 cm long and 2.0–3.5 mm wide. Lower bract usually 2.0–3.0 cm 

long, overtopping inflorescence. Inflorescence composed of one subsessile and 6–13 

ellipsoid-ovoid to ovoid-hemispherical 6–12-flowered pedunculate clusters. Peduncles 

straight, 2.0–3.5 cm long, densely papillose, often with secondary branches. Bractlets 

1.2–1.7 mm long, subentire. Tepals pale straw-brown, 2.4–3.1 mm long. Anthers 0.5–0.8 mm 

long, anther/filament length ratio 1.0–1.6; ovary 0.8–1.0 mm long; style 0.3–0.4 mm long; 

stigma 1.1–1.5 mm long. Capsule obovoid to subglobular, equalling perianth or shorter; capsule 

segments 1.9–2.1 mm long. Seeds ovoid to subglobular, 0.9–1.0 mm long, 0.75–0.8 mm 

wide; appendage 0.4 mm long. 2n=36 [12AL+24BL], fide J.Kirschner, Ann. Bot. Fenn. 29: 

238, 240 (1992). 

Known from Bulgaria and Greece. 13: BUL, GRC. Grassy mountain slopes, above 1000 m. 

Map 94. 

13. BULGARIA: Sofia, Gorni Lozen, 950–1050 m, 29 May 1988, J.Kirschner L110 (PRA); Sofia, Gorni 

Lozen, 950–1050 m, 17 Jun 1991, J.Kirschner L512, L513, L524 (PRA). GREECE: Serrai, Mt Vrondous, 

S.Snogerup 5052 (LD). 

59e. Luzula multiflora subsp. hibernica Kirschner & T.C.G.Rich, Watsonia 21: 92 (1996) 

T: Ireland, Mayo (Vice county H27, West Mayo), Cluddnaun (grid reference G/018.285), Jul 1989, 

T.C.G. Rich L254; holo: PR; iso: BEL, BM, DBN, E, K, LD, PRA, TCD. 

Illustration: J.Kirschner & T.C.G.Rich, Watsonia 21: 93, fig. 2 (1996) 

Stems 12–27 (–35) cm tall, usually slender. Basal leaves ±densely to subsparsely ciliate, 

2.2–2.8 (–3.5) mm wide. Cauline leaves usually 2–3. Upper cauline leaves 4.0–8.5 cm long, 

1.5–2.2 (–2.5) mm wide. Lower bract usually 1.7–4.3 (–5.5) cm long, shorter than or 

overtopping inflorescence, often reddish. Inflorescence with pedunculate clusters; peduncles 

mostly straight, some flexuose, usually at least one in a plant recurved, sometimes with 

secondary branches, smooth. Clusters 5–9 (–18); subsessile clusters 1–2, ellipsoidal-ovoid to 

ovoid, rarely hemispherical; flowers in pedunculate clusters 5–8 (–10); bractlets 1.5–1.8 mm 

long, ciliate. Tepals pale straw-brown or brown; outer sepals ±equalling inner ones or often 

up to 0.4–0.6 mm longer, 2.6–3.2 mm long, acuminate. Anthers 0.6–0.7 mm long, with 

anther-filament length ratio 1.0–1.2; ovary 0.9–1.0 mm long; style 0.6–0.7 mm long; stigma 

1.2–1.4 mm long. Capsules equalling perianth or shorter; capsule segments 1.9–2.2 mm long; 

seeds ovoid, 0.8–0.9 (–1.0) mm long, 0.7 mm wide; appendage 0.2–0.3 mm long. 2n=24 

[24AL], fide J.Kirschner, Folia Geobot. Phytotax. 30: 392 (1995). Fig. 25. 

Probably endemic in Ireland; in the west probably the only subspecies of L. multiflora. 

10: IRE. Mesotrophic grasslands along streams, track and road verges, wood clearings, 

usually at low altitudes. Map 95. 

10. IRELAND: North Kerry, Ross Castle, J.Kirschner, T.C.G.Rich & G.Kay (PRA); West Galway, Connemara, 

Kilmeelickin, J.Kirschner, T.C.G.Rich & G.Kay (PRA); West Mayo, Ballinglen, Keerglen R., J.Kirschner, 

T.C.G.Rich & G.Kay (PRA); Kerry, Killarney, Upper Lake, B.Welch 295 (UPS).


Figure 25. Luzula multiflora subsp. hibernica. A, habit; B, inflorescence; C, perianth and 

capsule; D, seed. (A–D, J.Kirschner & T.C.G.Rich L254, BM, PR). Scale bars: A, B = 1 cm; 

C, D = 1 mm. Drawn by A.Sier. Reproduced from Kirschner & Rich, Watsonia 21: 92, 

fig. 2 (1996). 

115

116 


There is evidence pointing to an alloploid origin of subsp. hibernica following hybridisation 

between L. pallescens and L. campestris; see J.Kirschner, Folia Geobot. Phytotax. 30: 

283–290 (1995). 

59f. Luzula multiflora subsp. monticola Kirschner, Pl. Syst. Evol. 200: 6 (1996) 

T: Spain, Huesca, Bisaurri, de la Fuente de la Muria a la Fabosa de la Muria, 1370–1540 m, 5 Jul 1988, Sesé et 

al., cultivated as L305; holo: PR; iso: PRA, JACA. 

Plants 13–26 cm tall. Basal leaves sparsely ciliate, 2–6 mm wide; cauline leaves 1 or 2, 

3.5–7.0 cm long, 2–3 mm wide. Lower bract c. 2.5–4.2 cm long, usually longer than 

inflorescence. Inflorescence composed of 3–8 usually short-peduncled narrowly ovoid 

c. 6–9-flowered clusters, at least partly subcongested. Peduncles ±straight, smooth, usually 

to 3.5 cm long (rarely to 8 cm). Tepals straw-brown to pale brown, (2.5–) 2.6–2.7 (–2.8) mm 

long. Anthers 0.6–0.7 mm long, anther/filament length ratio 1.2–1.4; ovary 0.7–0.8 mm long; 

style 0.3–0.4 mm long; stigmas 1.1–1.4 mm long. Capsule obovoid to subglobose, equalling 

to exceeding perianth; capsule segments 2.1–2.6 mm long. Seeds (0.8–) 0.9 mm long, 

(0.6–) 0.7 mm wide; appendage (0.3–) 0.4 mm long. 2n=24 [24BL], fide J.Kirschner, Pl. 

Syst. Evol. 200: 7 (1996). 

Known from the southern part of the Pyrenees in Spain. 12: SPA. Wet grasslands in the 

mountains. Map 96. 

12. SPAIN: Lérida, Sorpe, Puerto de la Bonaigua, 5 Aug 1986, P.Montserrat & L.Villar (JACA, PRA). 

Hybrids 

Luzula alpina Hoppe × L. multiflora (Ehrh.) Lej. 

A hybrid, sterile plant between a partially agmatoploid and a tetraploid parent 

(2n=28=20AL+8BL) was observed and collected in Austria [Hohe Tauern, Grossglockner, 

15 Jun 1953, M.Palm, cultivated by H.Nordenskiöld H1-1 (UPS)], and studied by 

H.Nordenskiöld (unpubl.). 

Luzula ×danica H.Nordensk. & Kirschner, Preslia 63: 109 (1991) = Luzula congesta 

(Thuill.) Lej. × L. multiflora (Ehrh.) Lej. 

T: Denmark, between Gram and Ribe, H.Nordenskiöld MT-3, holo: UPS. 

A hybrid known from Great Britain, Denmark and Sweden; seldom among parental plants. 

Fertile. 

Luzula campestris (L.) DC. × L. pallescens Sw. 

Hybrid plants rarely found between parents, up until now observed in Sweden only. Probably 

fertile. 

Luzula ×bogdanii Kirschner, Preslia 63: 105 (1991) = Luzula campestris (L.) DC. × L. fallax 

Kirschner 

T: Bulgaria, the Pirin Mtns, Rozhen, Vlashkija Put, J.Kirschner L117; holo: PR; iso: LD, PRA. 

At the places of contact between the parental species, the hybrid may form large populations. 

Up to now known from S Bulgaria. It is almost normally fertile and includes a number of 

hybrid karyotypes with mixed AL and BL chromosomes. 

Luzula ×media Kirschner, Preslia 63: 107 (1991) = Luzula divulgata Kirschner × 

L. multiflora (Ehrh.) Lej. 

T: [Czech Republic] Bohemia, Rakovník, Nový Mlýn, silva Brabečka, 16 Jun 1987, J.Hašková, J.Kirschner & 

F.Krahulec; holo: PR; iso: PRA


Found once between parents in the Czech Republic; documented karyologically [see 

J.Kirschner, Preslia 63: 107 (1991)]. At least partially fertile. 

Luzula ×heddae Kirschner, Preslia 63: 93 (1991) = Luzula campestris (L.) DC. × L. sudetica 

(Willd.) Schult. 

T: [Czech Republic] Bohemia, Hlinsko, Rváčov, 9 Jul 1990, L.Kirschnerová & J.Kirschner; holo: PR; iso: PRA. 

Found occasionally at the contact of parental species. Known from Sweden, Czech Republic, 

Switzerland and Spain, e.g., Sweden, Västmanland, Ramsberg, 29 Jun 1935, G.Samuelsson 

(LD); Dalsland, Edsleskog, 12 Jun 1918, G.Samuelsson (LD). A totally seed-sterile hybrid. 

Luzula ×hybrida H.Lindb. ex Kirschner, Preslia 63: 93 (1991) = Luzula pallescens Sw. × 

L. sudetica (Willd.) Schult. 

T: Finland, Savonia borealis, Jorois [Joroinen], Lapinmäki, 2 Jul 1904, H.Lindberg; holo: LD; iso: [Pl. Finl. 

Exs.] 158, e.g. BP, H, K, P, PRC, S 

A rare, sterile hybrid known from the Czech Republic, Sweden, Finland and N Russia, e.g., 

Sweden, Torne Lappmark, Vittangi, 28 Jul 1929, O.Holmberg (LD); Finland, Posio, Keski- 

Posio, Salmela, 4 Jul 1949, G.Marklund (H, LD). 

Doubtful taxa 

Luzula campestris var. major Bluff & Fingerh., Comp. Fl. German. 1: 444 (1825). 

T: [Germany, otherwise not designated] 

Luzula multiflora var. pallescens Bluff & Fingerh., Comp. Fl. German. 1: 444 (1825). 

T: [Germany, otherwise not designated] 

Luzula campestris var. groenlandica Laest., Bot. Not. 1858: 144 (1858). 

T: Grönland, J.Vahl; holo: n.v. [possible duplicate: Igalico, Aug 1828, J.Vahl (W)]. 

Luzula campestris var. subspicata Laest., Bot. Not. 1858: 144 (1858). 

T: [Sweden] Karesuando, L.L.Laestadius; holo: n.v. 

Luzula campestris [?] f. hornemannii Blytt, Norges Fl. 1: 297 (1861). 

T: 'Juncus congestus Fl. D. tab. 1692'; syn; [Norway] 'paa Tørvgrund i de lavere, isaer vestlige Kystegne', 

M.Blytt; syn: n.v. 

Luzula campestris a. [unranked] pseudonemorosa Schur, Enum. Pl. Transsilv. 682 (1866). 

T: [Romania] 'Götzenberg, Preschbe, oberhalb Resinar', F.Schur; syn: n.v. 

Luzula multiflora var. colchica V.I.Krecz., Žurn. Russk. Bot. Obshch. Akad. Nauk S.S.S.R. 

12: 490 (1928). 

T: not indicated. 

Luzula multiflora subsp. thianschanica V.I.Krecz., Žurn. Russk. Bot. Obshch. Akad. Nauk 

S.S.S.R. 12: 490 (1928). 

T: Tian Shan [Mtns], otherwise not designated. 

Luzula multiflora subsp. incrassata V.I.Krecz., Žurn. Russk. Bot. Obshch. Akad. Nauk 

S.S.S.R. 12: 490 (1928). 

T: Sakhalin, Okhotsk region, Kamchatka [otherwise not designated]. 

117

118 


Luzula multiflora var. gracilis Buchenau ex V.I.Krecz., Žurn. Russk. Bot. Obshch. Akad. 

Nauk S.S.S.R. 12: 490 (1928). 

T: Russian Far East, Ayan, Avacha R. basin [not located at LE]. 

Doubtful Hybrid 

Luzula ×winderiae Murr, Magyar Bot. Lapok 28: 67 (1930). 

T: [Austria] Vorarlberg, supra lacum Spullersee montis Arlberg, c. 1900 m, 7 Aug 1916, J.Murr; holo: n.v. 

The name was reported to belong to a hybrid between L. multiflora var. alpestris 

[=L. alpina] and L. spicata. The existence of the hybrid is very doubtful. 

PREDOMINANTLY AFRICAN SPECIES 

60. Luzula mannii (Buchenau) Kirschner & Cheek, Kew Bull. 55: 900 (2000) 

Luzula campestris var. mannii Buchenau, Bot. Jahrb. Syst. 12: 159 (1890); Luzula campestris subsp. mannii 

(Buchenau) Weim., Svensk Bot. Tidskr. 40: 173 (1946); isonyms: Sóo, Acta Bot. Acad. Sci. Hung. 16: 367 (1971); 

Fern.Casas, Fontqueria 33: 41 (1992). T: [Cameroon] Kamerun-Gebirge, 10000–13500 ft, Mann 2108; 

lecto: K; iso (fragment): W; Fernando Po, 8500', Mann 658, 1467; para K; Cameroons, H.H.Johnston 48 

[the latter erroneously cited as Mann 48 by F.G.P.Buchenau]; para: K. 

Perennials 25–60 cm tall; rhizome horizontal or ascending; stolons short, ascending. Leaves 

with obtuse tip and densely papillose-serrulate margins, hairy to densely hairy. Lower bract 

leaf-like, usually shorter than inflorescence. Inflorescence of mostly pedunculate clusters. 

Peduncles smooth, usually with short secondary branches. Capsule shorter than perianth. 

Seeds ovoid-subglobular, 0.6–0.8 mm wide; appendage almost invisible, less than 0.1 mm, 

hyaline. 

A rare endemic with three disjunct populations in tropical Africa (Equatorial Guinea, 

Cameroon and the Uganda/Kenya border). Two subspecies are recognized. 

Anther/filament length ratio c. 1.5; peduncles ±straight and rigid, 0.3–0.5 mm 

in diam.; tepals blackish castaneous brown 60a. subsp. mannii 

Anther/filament length ratio 0.3–0.6; peduncles flexuose to arcuate, 0.2–0.3 mm 

in diam.; tepals pale castaneous brown to dark brown, usually with a narrow 

pale margin and often with a pale yellowish-stramineous middle part 60b. subsp. gracilis 

60a. Luzula mannii (Buchenau) Kirschner & Cheek subsp. mannii 

Plants 25–60 cm tall. Basal leaves usually 7–9 (–13) cm long, (4–) 5–6 (–8) mm wide; 

cauline leaves 2–3, usually 3.5–7.0 cm long and 2.5–4.5 mm wide. Stem rigid, erect, c. 1.7 mm in 

diam. Lower bract 2.0–6.5 cm long. Inflorescence composed of 3–4 ±sessile and 

2–7 pedunculate clusters; peduncles usually with secondary branches and 1–3 sessile clusters 

on each branch; peduncles straight, rigid, erect and patent (very rarely one of them 

±nodding), 0.3–0.5 mm in diam., 2–3 (rarely up to 11) cm long; clusters ±echinate, 

±globular, 0.6–0.9 (–1.2) cm in diam., usually with more than 13 flowers. Bracteoles long 

acuminate, membranous, often with cupreous middle part, up to 3 mm long, equalling 

perianth or shorter, densely ciliate. Tepals long acuminate, blackish castaneous brown, 

subequal, sometimes one or two outermost ones distinctly longer than inner ones, 2.6–3.0 mm 

long. Anthers usually 0.8–1.2 mm long; filaments 0.5–0.9 mm long; style 0.5 mm long; 

stigmas 1.5–1.8 mm long. Capsule ±globular, rounded above with an indistinct mucro; 

capsule segments 1.8–2.0 mm long and c. 1.5 mm wide, dark brown. Seeds 0.9–1.1 mm long, 

0.7–0.8 mm wide. 2n=42, fide J.K.Morton, Opera Bot. 121: 159–172 (1993), n.v. Fig. 26.


Restricted to the Cameroons Mts. in Cameroon, and to Bioco in Equatorial Guinea. 23: CMN, 

EQG. Mountain grasslands, along tracks. Map 97. 

23: CAMEROON: ‘Region der Grassteppe am grossen Kamerunberg’, 3400–3700 m, 17 Oct 1906, 

Wederbauer 50 (WRSL); Cameroons Mts., N of Hut 2, 10500 ft, J.K.Morton K780 (K, S); Cameroons Mts., N 

of Hut 2, 10500 ft, J.K.Morton 6904 (K); Cameroons Mountain, ‘beyond the second ridge’, 10000 ft, 

T.D.Maitland 1248 (K); Cameroons Mt., ‘shallow volcanic soil, 11500 ft, Hinds C17 (K); ‘supra Beam pagum 

2850 m supra mare’, 21 Jul 1891, P.Dusén s.n. (S). EQUATORIAL GUINEA: Bioco, ‘pico Basilé’, 2950 m, 

Fernández Casas 10176a (K); Bioco, ‘pico Basilé’, 2870 m, Fernández Casas 11201 (K); Carvalho 2534 (K); 

Fernando Po [Bioco], Pico St. Isabel, 3000 m, E.Guinea 2868 (K). 

60b. Luzula mannii subsp. gracilis (S.Carter) Kirschner & Cheek, Kew Bull. 55: 901 (2000) 

Luzula campestris var. gracilis S.Carter, Kew Bull. 17: 179 (1963). T: Uganda, Mbale District, Mt Elgon, 

Jan 1918, Drummer 3545; holo: K; iso: BM, NBG, P. 

Illustration: S.Carter, in Fl. Trop. E. Afr., Juncaceae 9, fig. 2 (1966). 

Plants up to 65 cm tall. Basal leaves usually 6–15 cm long, 3–4 mm wide; cauline leaves 3–4, 

5.5–12 cm long, usually 2–4 mm wide. Lower bract 1.0–1.5 cm long, usually 0.7–1.0 mm 

wide. Inflorescence composed of 1–2 subsessile and (4–) 6–10 (rarely to 12) pedunculate 

clusters; peduncles often with secondary branches and 1–3 clusters on up to 1 cm long 

secondary peduncles; peduncles flexuose to arcuate, 0.2–0.3 mm in diam., 2.0–2.5 (–4.5) cm 

long, ±smooth to ±sparsely papillose. Clusters oblong-cylindrical, 0.5–0.8 cm long and up to 

0.5 cm wide, 5–15-flowered. Bracteoles lanceolate-ovate, with outer ones long acuminate, 

1.5–2.0 mm long, shorter than perianth, sparsely ciliate. Tepals equal, 2.3–2.8 (–3.0) mm 

long, narrowly lanceolate, acute to acuminate, pale castaneous brown to dark brown, usually 

with a narow pale margin and often with a pale yellowish-stramineous middle part. Anthers 

usually 0.3–0.6 mm long; filaments usually 0.9–1.1 mm long; style c. 0.4 mm long; stigmas 

less than 1 mm long. Capsule ±globular, obtuse, not mucronate; capsule segments castaneous 

brown, usually (1.4–)1.5–1.8 mm long and 1.1–1.2 mm wide. Seeds 0.8–1.0 mm long, 

0.6–0.8 mm wide. 2n=24, c.30, fide I.Hedberg & O.Hedberg, Bot. Not. 130: 11 (1977). 

Fig. 27. 

Restricted to Kenya/Uganda boundary range of the Mt. Elgon group. 25: KEN, UGA. 

Mountain grasslands and open woodlands, along paths. Map 98. 

25: KENYA: Mt. Elgon, E slope above Japata Estate, c. 3000 m, O.Hedberg 186 (K, S, UPS); Mt. Elgon, 

Mbere R. valley S of Koitcut, 3120 m, Gillett 18433 (K); Mt. Elgon, ridge W of Suam R., 3350 m, S.W.Bie 

66340 (UPS). UGANDA: Mt. Elgon, W slope above Butadiri, 3000 m, O.Hedberg 4450 (K, UPS); Mt. 

Elgon, by Sala stream, 10500 ft, Wood 126 (K); Mt. Elgon, Liebenberg 1704 (K); Mt. Elgon, Haines 4162 (K); 

Mt. Elgon, F.Rose 10145 & 10146 (K). 

61. Luzula abyssinica Parl., Fl. Ital. 2: 310 (1852) 

Luzula spicata var. simensis Hochst., in G.H.W.Schimper, It. Abyssin., no. 1154, (c. 1842); Luzula spicata var. 

erecta E.Mey., Linnaea 22: 415 (1849), nom. illeg.; Luzula macrotricha Steud., Syn. Pl. Glumac. 2: 294 (1855), 

nom. illeg.; Luzula abyssinica var. simensis (Hochst.) Buchenau, in H.G.A.Engler, Pflanzenr. (iv.36) 25: 40 

(1906), nom. inval. T: Ethiopia, Semien, Demerki, Schimper [It. Abyssin.] 1154; holo: FI; iso: K, P, S, PRC. 

Luzula volkensii Buchenau, Bot. Jahrb. Syst. 21: 192 (1896); Luzula abyssinica var. volkensii (Buchenau) 

Engl., Festschr. Aschers. 556 (1904). T: [Tanzania, Kilimanjaro] Südabfall der Mawensi-Spitze, 3900 m, 30 

Oct 1893, G.L.A.Volkens 1365; holo: B, destroyed; iso: K, W. 

Luzula abyssinica var. kilimandscharica Engl., Festschr. Aschers. 556 (1904). T: [Tanzania] Moshi, 

Kilimandscharo, A.Engler 1834; syn: B, probably destroyed; Moshi, Kilimandscharo, A.Engler 1755; syn: B, 

probably destroyed; Kilimandsharo, Kibosho, Uhlig 1110; syn: B, probably destroyed; isosyn: EA, n.v.; 

[Tanzania] Mt Meru, Uhlig 609; syn: B, probably destroyed; isosyn: EA, n.v. 

Illustration: K.A.Lye, in Fl. Ethiop. Eritr. 6: 390, fig. 211.3 (1997). 

Maps: H.Weimarck, Svensk Bot. Tidskr. 40: 173, fig. 14 (1946); H.Meusel, E.Jäger & E.Weinert, Vergl. 

Chorol. Zentraleur. Fl. 1: 88 (1965). 

Perennial c. (15–) 20–40 (–50) cm tall; rhizome thin, creeping to ascending; stolons usually 

absent. Stem erect, often rigid; basal leaves often numerous (few in shaded plants), flat to 

slightly canaliculate, up to 10–14 cm × 5–6 mm, densely hairy; leaf tip obtuse, swollen. Stem 

119

120 


Figure 26. Luzula mannii subsp. mannii. A, habit; B, tepals; C, capsule segment; D, flower 

with open capsule; E, seeds; F, pistil; G, flower with bracteoles; H, flower cluster; I, bracteole. 

(A–I, Maitland 1248, K). Scale bars: A = 1 cm; B–G = 1 mm; H = 2 mm; I = 1 mm. Drawn 

by C.Speight. Reproduced by kind permission of the Board of Trustees of the Royal Botanic 

Gardens Kew, from Kirschner & Cheek, Kew Bull. 55: 902, fig. 1 (2000).


Figure 27. Luzula mannii subsp. gracilis. A, habit; B, flower cluster; C, seed; D, flower; 

E, pistil. (A–E, Dummer 3545, K). Scale bars: A = 4 cm; B = 5 mm; C = 1 mm; D = 3 mm; 

E = 2 mm. Drawn by M.Grierson. Reproduced by kind permission of the Board of Trustees 

of the Royal Botanic Gardens Kew, from Fl. Trop. E. Afr., Juncaceae, p. 9, fig. 2 (1966). 

121

122 


leaves 2–3 (–4), usually 8–13 cm × 3.0–4.5 mm, hairy; leaf tip obtuse. Lower bract 

herbaceous, c. 6 cm × 1 mm, erecto-patent, exceeding inflorescence, with obtuse tip; upper 

bracts gradually membranous; middle and upper cluster bracts membranous with long acute 

tip. Inflorescence erect, usually congested and forming an elongated dense head 2–3 (–5) × 

0.7–1.0 cm, very often ±interrupted in lower part and lobulate, composed of 6 to many dense 

clusters of 8–20 flowers, lower cluster sometimes remote or on a long peduncle at the axil of 

the upper cauline leaf; peduncle smooth. Bracteoles 2.5–3.0 mm long, white-membranous, 

long fimbriate-ciliate (whole inflorescence has a hairy appearance). Tepals subequal, 

lanceolate, (2.4–) 2.8–3.2 mm long, entire or with a few teeth near the apex, acute; outer 

tepal long acuminate, castaneous brown, whitish bordered or unbordered. Stamens 6; anthers 

c. 0.6–0.9 mm long, slightly shorter or longer than 0.6–0.7 mm long filaments. Style 0.4–0.6 mm 

long; stigmas 1.5–2.5 mm long. Capsule ±obovoid, obtuse or very slightly subacuminate, 

castaneous brown, shorter than tepals; capsule segments 1.9–2.1 × 1.2–1.4 mm. Seeds 

±ellipsoidal, castaneous brown, 0.9–1.1 mm long, 0.6–0.7 mm wide, with a broad whitish 

caruncle 0.10–0.15 mm long. 2n=24 [(Bie 66129, O.Hedberg 1216, herbarium label records; 

I.Hedberg & O.Hedberg, Bot. Not. 130: 11 (1977)]. 

Eastern part of Tropical Africa, from E Zaire and Rwanda to Tanzania, Kenya and Ethiopia. 

23: RWA, ZAI. 24: ETH. 25: KEN, TAN, UGA. In mountain grasslands, on rocky slopes, 

and along paths. Map 99. 

23. RWANDA: Muhavura, 3350 m, 20 Feb 1972, P.H.Auquier 2610 (AAU). 24. ETHIOPIA: Prov. Simien, 

Geech Plateau, 3650 m, Heinonen 1009 (UPS); Arussi Prov., Galama Mts, Boraluco, 3750 m, O.Hedberg 4180 

(UPS); Shoa Prov., Mt Wuchacha, 19 Nov 1972, I.Friis et al. 1233 (C). 25. KENYA: Mt Kenya, NW 

slopes, 3100 m, 19 Aug 1948, O.Hedberg 1963 (S); Mt Elgon, 3400 m, 6 Jun 1920, Lindblom (S). 

TANZANIA: Morogoro, Uluguru Mtns, Lukwangule, 2400 m, 19 Feb 1933, Schliessen 3507 (S); Kilimanjaro, 

Peters Hut, 3800 m, O.Hedberg 1216 (UPS); Mt Meru, above Olkakola Estate, 3700 m, 1 Nov 1948, 

O.Hedberg 2427 (S). UGANDA: Ruwenzori, Bujuku Valley, near Bigo Camp, 3400 m, 22 Mar 1948, 

O.Hedberg 387 (S). 

A variable species. Small-flowered forms are sometimes found in populations. Plastic in size 

of vegetative parts (slender forms found in shady places). 

62. Luzula africana Drège ex Steud., Syn. Pl. Glumac. 2: 294 (1855) 

T: [South Africa] Katberg, J.F.Drège 3963; syn: BOL, K, PR, PRC, S. 

Illustration: A.A.Obermeyer, in A.A.Obermeyer, J.Lewis & R.B.Faden, Fl. S. Afr. 4/2: 90, fig. 26 (1985). 

Map: H.Weimarck, Svensk Bot. Tidskr. 40: 173, fig. 14 (1946). 

Perennials, c. 20–50 cm tall, densely caespitose; rhizome short, vertical; stolons absent. 

Basal leaves up to 10–15 cm long, 2.5–5.0 mm wide, ciliate; cauline leaves 2–3; upper 

leaves 2.5–6.0 cm long, 2–3 mm wide; all leaves with obtuse swollen tip, margins papilloseserrulate. 

Lower bracts 1–2, usually 1.0–2.5 cm long, obtuse to subacute, usually 

±overtopping inflorescence. Inflorescence congested in a dense many-flowered head c. 1–2 × 

1.0–1.2 cm. Bracteoles 1.5–2.0 mm long, ±ovate, ciliate-fimbriate. Tepals subequal, 

glabrous, straw-brown to brown with darker middle stripe and pale border, lanceolate, 

acuminate, 2.8–3.5 mm long. Stamens 6; anthers 0.8–1.1 mm long; filaments 0.5–0.7 mm 

long; style 0.7–0.8 mm long; stigmas 2.0–2.4 mm long. Capsule pale brown, obovate, 

indistinctly apicate-submucronate; capsule segments usually 2.2–2.4 × 1.6–1.7 mm. Seeds 

oblong-ovoid, pale brown, 0.7–0.9 (–1.0) mm long, 0.6–0.7 mm wide; appendage 0.2–0.3 mm 

long. 

Endemic to South Africa. 27: CPP, LES, NAT, OFS, TVL. In moist grasslands in the 

mountains. Map 100. 

27. CAPE PROVINCE: top of Katberg, J.Hutchinson 1617 (BOL); Ladysmith Div., Toverkop, Swartberg, 

E.Esterhuysen 26785 (BOL, K); Doornbosch, Kondeveldberge, E.G.H.Oliver 5214 (NBG). LESOTHO: 

Mont aux Sources, 9500 ft. Flanagan 2008 (BOL, K); Leribe, 1600–1900 m, A.Dieterlen 734 (K, SAM); Sani, 

Hodgson’s Peaks, 3000 m, O.Hilliard & B.L.Burtt 9676 (E, S). NATAL: Drakensberg, 2450 m, Phillipson 

661 (K). TRANSVAAL: Dullstroom, Verlorenvallei Farm, 2550 m, Drews 36 (K); Mooi R., 2000 m, 

F.R.Schlechter 3336 (BM).

Part 6. JUNCACEAE 1: Rostkovia to Luzula - Species Plantarum ...

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