JUNCACEAE (Juncus) - Species Plantarum Programme

JUNCACEAE (Juncus) 

172. Juncus elliottii Chapm., Fl. South. U.S. 494 (1860) 

T: Florida, A.W.Chapman s.n.; lecto: NY247634, designated here by S.E.Clemants; isolecto: GH29637. 

Juncus elliottii var. polyanthemus C.Mohr, Contr. U.S. Natl. Herb. 6: 435 (1901). T: Alabama, Mobile, 

C.Mohr s.n.; holo: UNA. 

Illustrations: H.A.Gleason, Ill. Fl. N. U.S. 1: 398 (1958); R.K.Godfrey & J.W.Wooten, Aquat. Wetl. Pl. SE 

U.S. 562, fig. 327 (1979). 

Map: R.E.Brooks & S.E.Clemants, Fl. N. Amer. 22: 252 (2000). 

Perennials, 30–90 cm tall, caespitose; rhizome much branched; older roots with ellipsoid to 

pyriform tuberiform distal thickenings. Cataphyll single, maroon to brown, lax, acute, often 

with a filiform blade to 5 mm long; basal leaves 1–3, 2–16 cm long; Cauline leaves 1–2, 

2–16 cm long, compressed, perfectly septate (septa visible externally but not prominent), 

unitubular; tip acute; auricles 0.5–2 mm long, rounded, scarious. Lower bract leaf-like, 

lanceolate, 1–4 cm long, shorter than the inflorescence, erect to ascending; other bracts 

stramineous, narrowly ovate, 0.5–1.3 cm long. Inflorescence decompound to 

supradecompound, usually loose, rarely subdense, of 40–100 (–200) heads, (2–) 4–10 (–16) cm 

tall, with ascending to spreading branches; heads semiglobose to obpyramidal, 2–10flowered, 

3–5 mm in diam. Tepals unequal, lanceolate, greenish stramineous, later 

stramineous, often suffused reddish, acuminate; inner tepals (1.8–) 2.4–2.8 mm long; outer 

tepals (2.2–) 2.6–2.9 mm long. Stamens 3, not exserted; anthers 0.3–0.7 mm long; filaments 

0.5–1.0 mm long; style 0.1–0.2 mm long; stigmas c. 0.6 mm long. Capsules unilocular, 

narrowly obpyriform to narrowly ovoid, abruptly narrowing in acute apex, 2.4–2.9 mm long, 

castaneous-brown, ±exceeding perianth. Seeds ellipsoid, apiculate, 0.4–0.6 × c. 0.2 mm, 

distinctly c. 13–16-striate, irregularly reticulate; appendages absent. 2n = 40, fide 

R.E.Brooks & S.E.Clemants, Fl. N. Amer. 22: 252 (2000). 

Confined to the SE United States. 75: NWJ. 77: TEX. 78: ALA, ARK, DEL, FLA, GEO, 

LOU, MSI, NCA, SCA, TEN, VRG. Wet sands, peaty sands, or peat, exposed shores of 

ponds and lakes, depressions in savannahs and flatwoods, moist to wet, much-disturbed 

clearings, roadsides, and ditches; 0–700 m. Map 365. 

77. TEXAS: Harris Co., N of Webster, D.S.Correll 32914 (NY). 78. FLORIDA: Apalachicola, 

A.W.Chapman [G.Engelmann, Herb. Junc. Bor.-Amer. Norm.] 54 (NY, PR); Lake Co., Eustis, G.V.Nash 338 

(PR, WU); Jacksonville, A.H.Curtiss 5417 (WU). TENNESSEE: Coffee Co., E of Tullahoma, H.K.Svenson 

10260 (BKL). VIRGINIA: Sussex Co., 6.5 km NW of Waverly, M.L.Fernald & B.Long 6139 (BKL, NY); 

Princess Anne Co., Rifle Range, S of Rudy Inlet, L.B.Smith & A.R.Hodgdon [Pl. Ex. Gray.] 643 (NY). 

173. Juncus mertensianus Bong., Mém. Acad. St. Pétersbourg 6(2): 167 (1833) 

T: [Alaska, Sitka] Sitcha, K.H.Mertens s.n.; holo: LE; iso: PRC. 

Juncus ensifolius var. minor E.Mey., Linnaea 3: 373 (1828). T: Unalaska, Wickstroem s.n.; holo: n.v. 

Juncus mertensianus var. filifolius Suksd., Deutsche Bot. Monatsschr. 19: 92 (1901). T: Butterfly Lake, 

Skamania Co, Washington, 11 Jun 1891, W.N.Suksdorf 2042; holo: WS; iso: NY247682. 

Juncus duranii Ewan, Rhodora 47: 186 (1945); Juncus mertensianus var. duranii (Ewan) F.J.Herm., Leafl. W. 

Bot. 10: 85 (1964). T: California, Los Angeles Co, San Gabriel Mts., Mt Hawkins, Lilly Spring, J.A.Ewan 

10060; holo: COLO. 

Juncus slwookoorum S.B.Young, Rhodora 72: 486 (1970). T: Alaska, St. Lawrence Island, north shore of 

Boxer Bay Lake, S.B.Young 701; holo: GH29676; iso: GH29675. 

Illustrations: F.Buchenau, in H.G.A.Engler, Pflanzenr. (iv.36) 25: 201, fig. 96 (1906); W.L.Jepson, Fl. Calif. 

1: 253, fig. 42f, g (1921); C.L.Hitchcock & al., Vasc. Pl. Pacif. Northw. 1: 202 (1969); R.E.Brooks & 

S.E.Clemants, Fl. N. Amer. 22: 253 (2000). 

Maps: E.Hultén, Fl. Alaska 288 (1968); R.E.Brooks & S.E.Clemants, Fl. N. Amer. 22: 252 (2000). 

Perennials, 5–40 cm tall, rhizomatous, loosely caespitose. Rhizomes short-creeping to longcreeping, 

occasionally stoloniferous, 1–2 mm in diam., not swollen, with internodes to c. 1.5 cm 

or abbreviated. Cataphylls 1 or absent, stramineous to castaneous-brown, acute, with distinct 

scarious margins; blade short, thicker acicular. Basal leaves 0–1, 3–15 cm. Cauline leaves 1–3, 

rarely absent, 3–15 cm long, with upper 1–2 often reaching inflorescence and uppermost leaf 

sometimes suberect to erecto-patent seemingly as a stem continuation, terete, perfectly 

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SPECIES PLANTARUM — FLORA OF THE WORLD (2002) 

septate (septa not very distinct externally), unitubular; tip subulate; sheaths lax, with broad 

margins; auricles 1.0–1.2 mm long, rounded to acute, membranous or scarious. Lower bract 

leaf-like, green to castaneous, lanceolate to terete, 1–2.4 cm long, shorter than to longer than 

the inflorescence, erect; other bracts castaneous, lanceolate, 2.5–3 mm long. Inflorescence a 

single head (rarely a cluster of two heads), 0.5–1.6 cm long; heads spheric (to semiglobose), 

12–60-flowered, 4.5–15 mm in diam. Tepals unequal to subequal, lanceolate to lanceolateovate, 

dark purplish brown to black with indistinct dark castaneous margins, acute, with 

mucro subulate; inner tepals 2.3–4.3 mm long; outer tepals 2.4–4.9 mm long. Stamens 6, not 

exserted; anthers (0.6–) 1.1–1.7 mm long; filaments 0.6 (–1.7) mm long; style 0.5–0.7 mm 

long; stigmas 1.0–1.5 mm long. Capsules unilocular, obovoid, 1.9–3.5 mm long, obtuse or 

rounded to truncate, with mucro c. 0.2 mm long, castaneous-brown, equalling or shorter than 

perianth. Seeds narrowly ellipsoid, 0.5–0.6 × 0.2–0.25 mm, apiculate, pale brown, distinctly 

striate and reticulate; appendages absent or minute, whitish. 2n = 80, fide L.Hämet-Ahti, 

Ann. Bot. Fenn. 8: 156 (1971); 2n = 40, fide R.L.Taylor & G.A.Mulligan, Fl. Queen 

Charlotte Is. 2: 44 (1968). Fig. 104. 

Widely distributed along the Pacific coast of North America from California to Alaska and 

the Aleutian Is., often also in the Rocky Mtns from N Arizona and N New Mexico to 

Montana, Wyoming and Colorado in the northeast. 70: ASK, YUK. 71: ABT, BRC, SAS. 

73: COL, IDA, MNT, ORE, WAS, WYO. 76: ARI, CAL, NEV, UTA. 77: NWM. Montane to 

alpine meadows, stream banks, lake margins, and conifer woods; (400–) 1900–3300 m. 

Map 366. 

70. ALASKA: Kodiak Island, Old Harbour, W.J.Eyerdam 712 (NY). YUKON: Itsi Range, J.A.Calder & 

I.Kukkonen 27653 (NY). 71. ALBERTA: Waterton Lakes National Park, Cameron Lake, A.J.Breitung 

16161 (NY). BRITISH COLUMBIA: Tats Lake, L.E.Pavlick 83–162 (PRA); Vancouver Is., Strathcona 

Provincial Park, Marble Meadows, A.Ceska & O.Ceska 4403 (PRA). 73. IDAHO: Lemhi Co., S end of Big 

Clear Lake, D.Atwood 14036 (NY). OREGON: Wallowa Co., Wallowa National Forest, M.F.Denton 3323 

(NY). WYOMING: Fremont Co., Wind River Range, R.W.Scott 802 (NY). 76. CALIFORNIA: E slope, 

Mono Pass, H.N.Bolander [herb. Norm. 92]. NEVADA: Elko Co., Mountain City, Jack Ck Summit, B.Ertter 

4285 (NY, PRA). UTAH: Alta, M.E.Jones 1094 (NY). 

Populations from southern California with brown tepals; anthers equalling filaments, and 

rounded to acute translucent auricles have been separated as J. duranii, but the typical form 

is so highly variable that it can easily accommodate this local form. This species passes into 

J. nevadensis and has often been combined with that species. The two species can generally 

be separated and we are following recent treatments. 

174. Juncus chlorocephalus Engelm., Trans. Acad. Sci. St. Louis 2: 485 (1868) 

T: 'higher mountains of California', W.B.Hillebrand 2338; syn: MO, n.v.; Mount Dana, 10,000 ft, W.H.Brewer 

1804; syn: MO, n.v., GH; Yosemite Valley, 4000 ft., H.N.Bolander 6033 [G.Engelmann, Herb. Junc. Bor.- 

Amer. Norm.] 99; syn: BKL, MO, NY, PR; Carson City, Nevada. C.L.Anderson; syn: MO, n.v. 

Illustrations: W.L.Jepson, Fl. Calif. 1: 254, fig. 43, h, i (1921); L.Abrams, Ill. Fl. Pacific States 1: 363, fig. 

889 (1923). 


Perennials, 20–45 cm tall, caespitose; rhizome ±horizontal, short-creeping, very short-noded. 

Cataphyll single, stramineous or maroon, acute; basal leaves 1–2, 2–16 cm tall. Cauline 

leaves 2–3, 2–16 cm long, terete, perfectly septate, unitubular, to c. 1.3 mm in diam.; tip 

subulate; auricles 2–3.5 mm long, rounded, scarious. Lower bract ±scarious, distinctly 

3-veined, white or stramineous, narrowly ovate to linear-lanceolate, 1–1.8 cm long, shorter 

than to slightly exceeding inflorescence, erect or spreading; other bracts white or 

stramineous, ovate, 2.5–3 mm long. Inflorescence a single head (rarely a cluster of 2–3 

heads), 0.5–2 cm long; heads spherical, 15–25-flowered, 11–14 mm in diam. Tepals equal, 

lanceolate, light green to light pink or white, obtuse to ±mucronate, 3.1–4.2 mm long; midrib 

often prominent. Stamens 6, not exserted; anthers 1.6–2.2 mm long; filaments 0.7–1.5 mm 

long; style (1.7–) 2–3 mm long; stigmas 1.5–2.5 mm long. Capsules unilocular, broadly 

obovoid to ovoid, obtuse, 2.2–2.5 mm long, stramineous, shorter than perianth. Seeds 0.5–0.6 

× 0.2–0.3 mm; appendages absent.


Figure 104. Juncus nevadensis. A, habits of various forms; B, flowers of various forms; 

C, capsule; D, flower of the form correspondig to the type; E, seed; F, auricle. Juncus 

mertensianus. G, habit; H, auricle; I, flower with capsule; J, capsule; K, seed. Scale bars: 

A, G = 5 cm; B, C, D, I, J = 2 mm; F, H = 5 mm; E, K = 0.25 mm. Drawn by J.R.Janish. 

Reproduced with permission, from C.L.Hitchcock, A.Cronquist, M.Ownbey & J.W.Thompson, 

Vasc. Pl. Pacif. Northw. 1: 188, 202 (1969). © University of Washington Press. 

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Confined to the mountains of SW United States. 76: ARI, CAL, NEV. Sandbars, meadows, 

damp soil in rock outcrops and talus; 1400–3300 m. Map 367. 

76. CALIFORNIA: Mariposa Co., Yosemite National Park, Yosemite Ck, Tioga Rd, W.J.Dress 3577 (W); 

Mariposa Co., 6 km W of Lake Tenaya, Cooke, D.J.Pinkava, D.J.Keil & Weber 2130 (W); Placer Co., above 

Donner Pass, A.A.Heller 7178 (BKL, NY); Tuolumne Co., 17.5 km NE of Pinecrest, B.Ertter & A.Carter 7242 (NY); 

Eldorado, Desolation Valley, L.R.Abrams 12736 (BRNU). NEVADA: Douglas Co., Carson Range S of 

Spooner Summit, A.Tiehm 12033 (NY). 

175. Juncus nevadensis S.Watson, Proc. Amer. Acad. 14: 303 (1879) 

Juncus phaeocephalus var. gracilis Engelm., Trans. Acad. Sci. St. Louis 2: 484 (1868); Juncus mertensianus 

subsp. gracilis (Engelm.) F.J.Herm., Leafl. W. Bot. 10: 85 (1964). T: numerous syntypes, incl. Big Tree 

Grove, H.N.Bolander; syn: MO, n.v.; Tuolumne R., Headwaters, H.N.Bolander [G.Engelmann, Herb. Junc. 

Bor.-Amer. Norm.] 98; syn: MO, PR. 

Juncus suksdorfii Rydb., Bull. Torrey Bot. Club 26: 541 (1899); Juncus mertensianus var. suksdorfii (Rydb.) 

F.J.Herm., Leafl. W. Bot. 10: 86 (1964). T: Washington, Fulda, Falcon Valley, W.N.Suksdorf 217; syn: US, 

GH29682; W.N.Suksdorf 680; syn: GH; Spokane Co., Spangle, W.N.Suksdorf 464; syn: GH. 

Juncus badius Suksd., Deutsche Bot. Monatsschr. 19: 92 (1901); Juncus mertensianus var. badius (Suksd.) 

F.J.Herm., Leafl. W. Bot. 10: 86 (1964); Juncus nevadensis var. badius (Suksd.) C.L.Hitchc., in C.L.Hitchcock 

& al., Vasc. Pl. Pacif. Northw. 1: 201 (1969). T: Washington, Klickitat Co, Falcon Valley, 25 Jun 1892, 

W.N.Suksdorf 2144; holo: US; iso: GH29606, NY247565. 

Juncus columbianus Coville, Proc. Biol. Soc. Wash. 14: 87 (1901); Juncus nevadensis var. columbianus 

H.St.John, Fl. SE. Washington 82 (1937); Juncus mertensianus var. columbianus (Coville) F.J.Herm., Leafl. 

W. Bot. 10: 86 (1964). T: Washington, Whitman Co, Pullman, A.D.E.Elmer 235; holo: US; iso: BKL, 

NY247608. 

Juncus truncatus Rydb., Bull. Torrey Bot. Club 31: 399 (1904). T: Colorado, Meadow Height, 1898, 

C.L.Shear & E.A.Bessey 4323; holo: NY247742. 

Juncus inventus L.F.Hend., Rhodora 32: 21 (1929); Juncus nevadensis var. inventus (L.F.Hend.) C.L.Hitchc., 

in C.L.Hitchcock & al., Vasc. Pl. Pacif. Northw. 1: 201 (1969). T: Oregon, Lane Co., Siltcoos Lake, 

2 miles [3.2 km] from the ocean, L.F.Henderson 6091; holo: ORE96223. 

Illustrations: F.Buchenau, in H.G.A.Engler, Pflanzenr. (iv.36) 25: 203, fig. 97, 205, fig. 98 (1906); C.L.Hitchcock & 

al., Vasc. Pl. Pacif. Northw. 1: 202 (1969); R.E.Brooks & S.E.Clemants, Fl. N. Amer. 22: 253 (2000). 


Perennials, 5–70 cm tall, rhizomatous, loosely caespitose. Rhizome stoloniferous, 1 mm in diam., 

not swollen, often densely branched. Cataphylls 1 or absent, maroon or stramineous, acute; basal 

leaves 1–3, 1.5–31 cm long; Cauline leaves 1–2, 1.5–31 cm long, laterally flattened to subterete, 

perfectly septate (septa visible in dry condition), unitubular, 0.5–1.5 (–2.0) mm wide; tip acute 

to acuminate; sheaths broadly bordered; auricles scarious, 1–3.2 mm long, rounded to acute. 

Lower bract leaf-like, linear, 1.5 cm long, shorter than the inflorescence, erect; other bracts 

reddish-green, lanceolate, 6–6.5 mm long. Inflorescence umbelloid to anthelate, of (1–) 4–8 (–11) 

heads, 2–18 cm tall, with erect to spreading branches; heads semiglobose to obpyramidal 

(rarely subspheric), 3–11-flowered, 5–14 mm in diam. Tepals subequal, lanceolate to broadly 

lanceolate, acute to acuminate, with tip subulate, stramineous-brown to dark castaneousbrown, 

rarely pale stramineous; outer tepals slightly longer, 2.8–6.2 mm; inner tepals 2.4–6 mm 

long; margins broad, membranous to scarious; central band greenish. Stamens 6, not exserted; 

anthers 1.0–2.0 mm long; filaments 0.4–0.7 mm long; style (0.7–) 1.0–1.8 mm long; stigmas 

1.2–2.5 mm long. Capsules unilocular, ellipsoid, trigonous, acute below a c. 0.5 mm rostrum, 

2.3–3.7 mm long, castaneous-brown, shorter than to slightly exceeding perianth. Seeds ovoidellipsoid, 

apiculate, 0.4–0.5 × 0.2 mm, brown, distinctly reticulate; appendages absent. Fig. 104. 

In western North America, centred in the Rocky Mtns and reaching S Alberta and 

S Saskatchewan in the north. 71: ABT, SAS. 73: COL, IDA, MNT, ORE, WAS, WYO. 

76: ARI, CAL, NEV, UTA. 77: NWM. Wet banks along the margins of streams and lakes, 

marshes, bogs, sloughs, and wet or boggy prairies; 0–2600 m. Map 368. 

73. IDAHO: Camas Co., S edge of Macon Flat, B.Ertter 2413 (NY). OREGON: Crook Co., Black Butte, 

W.C.Cusick 2681 (PR). WASHINGTON: Clallam Co., Lake Ozette, J.W.Thompson 9418 (BKL); Klickitat 

Co., Bingen, W.N.Suksdorf 5055 (NY); Klickitat Co., Falcon Valley, 1907, W.N.Suksdorf [A.Kneucker, Cyper. 

Junc. Exs.] 205 (PR, PRA). 76. CALIFORNIA: Douglas Co., Ridges Ranch, J.M.Johnson 107 (NY).


NEVADA: Douglas Co., Pine Nut Mountains, A.Tiehm 12227 (NY, UTA); Sierras, Tuolumne R., Headwaters, 

H.N.Bolander [G.Engelmann, Herb. Junc. Bor.-Amer. Norm.] 98 (MO, PR). 

A variable species that has been broken down into five separate taxa in the past; more 

recently, most of the variants have not been separated. The Oregon coastal form 

[J. nevadensis var. inventus (L.F.Hend.) C.L.Hitchc.] is at one extreme of the variation 

having only a single head, fairly dark brown tepals 3.5–4.5 mm long; anthers only slightly 

longer than the filaments and a distinct habitat. However, the J. mertensianus var. 

columbianus segregate approaches var. inventus in number of heads and the other characters 

are so highly variable in the complex they can not be used alone to separate this variety. 

Therefore, infraspecific taxa are not recognized at this time. 

176. Juncus dubius Engelm., Trans. Acad. Sci. St. Louis 2: 459 (1868) 

T: California, Mariposa, Clark's Meadow near Big Tree Grove, H.N.Bolander 6032 [G.Engelmann, Herb. 

Junc. Bor.-Amer. Norm.] 52; syn: MO, NY, PR. 

Juncus rugulosus Engelm., Bot. Gaz. 6: 224 (1881); Juncus dubius f. rugulosus (Engelm.) Hoover, Leafl. W. Bot. 10: 

340 (1966). T: California, San Bernardino Mts.; Castac Lake. W.G.Wright s.n.; syn: MO, GH29674. 

Illustrations: W.L.Jepson, Fl. Calif. 1: 256, fig. 44a–e (1921); L.Abrams, Ill. Fl. Pacific States 1: 365, fig. 

893, 894 (1923). 


Perennials, 30–75 cm tall, rhizomatous, forming loose stands. Rhizome short-creeping to 

long-creeping, 2–3 mm in diam., pale brown, not swollen. Stem usually densely papilloserugulose, 

rarely ±smooth. Cataphylls 1–2, pink to stramineous, acute; basal leaves 1–2, 20–30 cm 

long. Cauline leaves 1–2, 20–30 cm long, terete, perfectly septate (septa visible externally, 

prominent in dry condition), rugulose or rarely smooth, unitubular; tip acuminate; sheaths 

broadly bordered; auricles 1–4.9 mm long, rounded, membranous. Lower bract herbaceous, 

stramineous, lanceolate to linear, 1.2–3.2 cm long, shorter than the inflorescence, erect; other 

bracts stramineous, lanceolate, 4–6 mm long. Inflorescence diffuse, decompound, of 25–66 

heads, usually 7–15 cm long, with main branches erect to erecto-patent and other branches 

often ±patent; heads semiglobose to obovoid, 6–10-flowered, 5–10 mm in diam. Tepals 

subequal, lanceolate, stramineous to brown, acuminate; outer tepals carinate, with prominent 

midrib, (2–) 2.5–3.4 mm long, stramineous-brown to castaneous, acuminate, with margins 

broad and membranous; inner tepals often slightly longer, flat, subacute to mucronate, 

(2–) 2.6–3.6 mm long, distally with broad membranous margins. Stamens 6, not exserted; 

anthers 0.6–1.0 mm long, c. 1.5 times as long as 0.5–0.7 mm filaments; style 0.5–0.6 mm 

long; stigmas 0.6–1.1 mm long. Capsules unilocular, tapering to a subulate tip from narrowly 

ovoid base, trigonous, (2.4–) 3–3.9 mm long, castaneous-brown, ±exceeding perianth. Seeds 

ellipsoid-ovoid, apiculate, 0.3–0.5 × 0.2–0.3 mm, pale brown, distinctly reticulate; 

appendages absent. 2n = 40, fide N.A.Harriman & D.Redmond, Rhodora 78: 735 (1976). 

Restricted to California and northernmost NW Mexico. 76: CAL. 79: MXN. Montane 

meadows, river beds, canyons, arroyos; 100–1600 m. Map 369. 

76. CALIFORNIA: San Bernardino Co., San Gabriel Mtns, Cajon Ck, R.F.Thorne, C.W.Tilforth & al. 42164 

(NSW); San Bernardino Mtns, S.B.Parish & W.F.Parish 1026 (PR); Kern Co., 3.2 km above Kernville, 

J.T.Howell 44112 (NY); San Diego Co., Cleveland National Forest, E slope Aqua Tibia Mt., D.L.Banks & 

S.Boyd 5 (NY); Sonoma Co., Laguna de Santa Rosa, R.Rubtzoff 4490 (NY); Tulare Co., Kaweah River at Three 

Rivers, J.T.Howell 50286 (NY); Los Angeles Co., Buckhorn Station, R.F.Thorne & L.E.DeBuhr 42239 (NSW). 

This species has rugulose stems and leaves throughout most of its range, but on the periphery 

of its range (in Mariposa, San Diego, and Sonoma Counties, California) the plants are smooth. 

177. Juncus wallichianus J.Gay ex Laharpe, Essai Monogr. Jonc. 51 (1825) 

T: Nepal, 1821, N.Wallich; syn: G, W. 

Juncus indicus Royle ex D.Don, Proc. Linn. Soc. London 1: 10 (1839) & Trans. Linn. Soc. London 18: 323 

(1840). T: [in 1839 not indicated]; Nepal [Kathmandu], N.Wallich 8999; syn: G-DC, K, W. 

Juncus monticola Steud., Syn. Pl. Glumac. 2: 301 (1855). T: [India] in montibus Nilagiri, R.F.Hohenacker [Pl. 

Ind. Orient.] 951; lecto: P, fide K.L.Wilson & L.A.S.Johnson, Telopea 9: 364 (2001); iso: E, K, L, P, PR, W. 

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Juncus prismatocarpus subvar. unitubulosus Buchenau, Bot. Jahrb. Syst. 12: 311 (1890); Juncus leschenaultii 

var. unitubulosus (Buchenau) Novikov, Nov. Sist. Vyssh. Rast. 19: 59 (1982). T: numerous syntypes from 

India and Japan, incl. Japan, Hiogo, Wawra 1453 (W); Nilgherries, Perrotet 1202 (W); East Benghal, 

W.Griffith 5459 p.p. (W); Nepal, 1821, N.Wallich (G-DC, W). 

Juncus koidzumii Satake, J. Jap. Bot. 12: 89 (1936). T: Japan, Honsyu, prov. Settu, Hyogo Prefecture, circa 

Takarazuka, 4 Aug 1934, N.Ui; holo: TI; iso: K. 

Juncus nikkoensis var. pinifolius Satake, J. Jap. Bot. 12 (8): 577 (1936). T: [Japan] Kyushu, Prov. Bungo, 

Mt Kuzyuzan, 22 Sep 1922, Z.Tashiro; holo: TI. 

Juncus pseudokrameri Satake, Rep. Sci. Exped. Manchoukuo, sect. 4 [=Index Fl. Jehol.] 4: 107 (1936). T: China, 

Manchuria, Prov. Hsing-an occid., prope Chih-feng, 26 Sep 1933, T.Nakai, M.Honda & M.Kitagawa; holo: TI. 

Juncus ohwianus Kao, in H.L.Li, Fl. Taiwan 5: 150 (1978). T: [citation: Hsinchu: Supachian, Simada 1027] 

Taiwan, Sintiki (18 sen[?]),12 Aug 1923, Y.Simada 1027A; holo: TAI. 

Juncus prismatocarpus subsp. teretifolius K.F.Wu, Acta Phytotax. Sin. 32: 456 (1994). T: [citation: 

Guangdong, Conghua, Sanjiao Shan, 31 May 1932, W.T.Tsang 20636] China, Kwangtung [Guandong], Sam 

Kok Shan, Tsungfa-Lungmoon District, 31 May 1932, W.T.Tsang 20636; holo: ISBC; iso: K, P [the 

transliteration of the Chinese ideograms for the locality is very different between 1932 and 1994, but the 

ideograms themselves are the same on the label and in the paper]. 

Juncus auritus K.F.Wu, Acta Phytotax. Sin. 32: 453 (1994). T: China, Yunnan, 1935, Chi-wu Wang 63454; 

holo: NAS, n.v. 

Juncus yanshanuensis Novikov, Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 103(4): 70 (1998); Juncus 

sphaerocephalus K.F.Wu, Acta Phytotax. Sin. 32: 453 (1994), nom. illeg. T: China, Yunnan, Yanshan, 

Shihtze Shan, 1200 m, 9 Nov 1939, Chiwu Wang & Ying Liu; holo: PE. 

Illustrations: Wu Kuofang, Fl. R. P. Sinicae 13(3): 187, tab. 43 (1997); Muh-Tsuen Kao & C.E.Devol, in Hui 

Lin Li & al., Fl. Taiwan 5: 151, tab. 1303 (1978), as J. ohwianus; K.L.Wilson & L.A.S.Johnson, Telopea 9: 

359, fig. 1 (2001). 

Perennials, 30–60 cm tall, usually densely caespitose, rarely shortly rhizomatous; stem 

terete, 1.0–1.8 mm in diam. Cataphylls 0–2. Cauline leaves 2–3, spread along stem and 

±equalling it, unitubular, perfectly septate, terete, 1.0–1.4 mm in diam.; auricles scarious, 

usually 1.0–2.0 mm long, ±acute. Lower bract leaf-like, 2.5–7 cm long, shorter than 

inflorescence. Inflorescence terminal, 1–12 cm long, decompound, diffuse, of 6–20 mostly 

semiglobose, (4–) 10–25 (–40)-flowered clusters. Tepals ±equal, narrowly lanceolate, 

(2.2–) 2.5–3.3 (–3.5) mm long, acuminate, with tips sometimes slightly recurved at maturity, 

greenish, later stramineous-brown (often red-tinged at apex); margins membranous, 0.1–0.2 mm 

wide. Stamens 3, shorter than perianth; anthers 0.4–0.7 mm long; filaments c. 0.8–1.2 mm 

long; style c. 0.1–0.2 mm long; stigmas 0.5–1.0 mm long. Capsule unilocular, narrowly 

ovoid to narrow-ellipsoid, c. 3.0–4.2 mm long, acute to broad-acute, with a short rostrum 

0.2–0.5 mm long, usually exceeding perianth, golden brown to castaneous-brown. Seeds 

ellipsoid, apiculate, 0.5–0.6 × c. 0.2 mm, pale brown, longitudinally c. 20-striate, irregularly 

transversally reticulate, areas finely lineolate; appendages absent. Fig. 90. 

Widely distributed in subtropical and tropical Asia, from Nepal, India and Sri Lanka in the 

west to Japan and Malesia and Papuasia in the east and southeast. Recorded from Sakhalin or 

Kuril Is. but material not seen. Introduced in Mauritius. 29: mau. 31: SAK, PRM. 36: CHC, 

CHH, CHI, CHM, CHN, CHS. 38: JAP, KOR, TAI. 40: ASS, BAN, EHM, IND, NEP, SRL. 

41: LAO, MYA, THA, VIE. 42: JAW, SUM. 43: NWG. Swamps, wet ditches, along streams. 

Map 370. 

29. MAURITIUS: Ile Maurice, Beau Bassin, 3 Apr 1974, J.Bosser 21840 (P). 36. CHINA SOUTHEAST: 

Hong-Kong, Fort Victoria, 20 Apr 1895, E.Bodinier 1113 (W); Canton [Guangzhou], 3 Apr 1884, T.Sampson 

(K). 38. JAPAN: Kiusiu, Simobara Province, Oyo lake, 1863, C.J.Maximowicz (K). 40. ASSAM: Khasia, 

W.Griffith s.n. (K). INDIA: S India, High Wavy Mtn, E.Blatter & R.Hallberg 328 (K). NEPAL: 

Gurjakhani, 2600 m, J.D.A.Stainton, W.R.Sykes & R.H.J.Williams 3681 (BM, P). SRI LANKA: Sri Lanka, 

Uva, Badulla Distr., Farr Inn, 21 Jul 1970, 2300 m, D.Clayton 5518 (K). 42. JAVA: Priangan, 2450 m, C.G. van 

Steenis 4090 (K). 43. PAPUA NEW GUINEA: West Papua, Lake Habbema, L.Brass 9046 (BRI, CANB, K, L)


178. Juncus leptospermus Buchenau, Bot. Jahrb. Syst. 6: 203 (1885) 

T: [India] Assam (Khasia), am unteren Brahmaputra, 4–6000 Fuss. J.D.Hooker & J.Thomson; syn: K; Assam, 

Khasia, 4–5500', C.B.Clarke; syn: K, several specimens; East Bengal, W.Griffith 5455; syn: K, W.Griffith 

5459 p.p.; syn: C, K; Maophlang, 5500', Khasia, 19 Oct 1872, C.B.Clarke 18687; syn: K. 

Juncus mairei H.Lév., Fedde Repert. Spec. Nov. Regni Veg. 11: 493 (1913). T: China, Yunnan, Tong 

Tchouan, Oct 1910, E.E.Maire 7357; holo: P. 

Illustrations: F.Buchenau, in H.G.A.Engler, Pflanzenr. (iv.36) 25: 197, fig. 95 (1906); Wu Kuofang, Fl. R. P. 

Sinicae 13(3): 187, tab. 43 (1997). 

Perennials, 37–85 cm tall, densely caespitose; rhizome short; stolons absent. Cataphylls 1–2, 

greenish to brownish, with broad membranous margins. Leaves 2–5, evenly spread out along 

stem, unitubular, perfectly septate, c. 3–20 cm long, usually 1.0–1.5 mm wide; sheaths slightly 

inflated; auricles obtuse to acute, 1–2 mm long, membranous or pale brownish. Lower bract 

stiff, erect, leaf-like, c. 1–4.5 cm long, shorter than inflorescence. Inflorescence densely 

branched, with 1–3 main branches, of 8–15 (–37) semiglobose to obconical, (2–) 4–10 (–19)flowered 

heads on short peduncles; middle bracts stramineous, aristate; capitulum bracts 

stramineous. Tepals subequal or inner ones longer, ±lanceolate, 2.4–3.0 mm long, pale brown, 

with distinct membranous margins. Stamens 3; anthers 0.4–0.6 mm long, pale yellow; 

filaments flattened, c. 1.2–1.7 mm long; style to 0.2 mm long; stigmas spreading, 0.5–1.0 mm 

long. Capsule unilocular, distinctly trigonous and narrowly oblong-lanceolate in outline, 

c. 3.8–4.6 mm long including a 0.2–0.5 (–0.7) mm rostrum, castaneous, shining, conspicuously 

exceeding perianth. Seeds narrowly ellipsoid, apiculate, c. 0.6–0.7 × c. 0.2 mm, pale brown to 

brown, longitudinally c. 18–24-striate, transversally reticulate; appendages ±absent. 

Scattered from Bhutan and Assam to C China. Alsop recorded from Guizhou but material not 

seen. Reports from other parts of China (e.g., Heilongjiang) are doubtful. 36: CHC, CHN. 

40: ASS, EHM. Wet grassy slopes, shores of ponds. Map 371. 

36. CHINA SOUTH-CENTRAL: Yunnan, Zhongdian Co., E of Zhongdian, 3340 m, [RBGE Exped. 1995] 379 (E); 

Zhongdian Co., W end of Bita Hai lake, 3560 m, [RBGE Exped. 1995] 410 (E); Zhongdian, 3240 m, [CLDE] 

258 (K); Yulong Shan, above the Lan gon Lake, 3200 m, [CLDE] 1140 (K). 

179. Juncus krameri Franch. & Sav., Enum. Pl. Jap. 2: 99 (1879) 

T: Japan, circa Simoda, P.A.L.Savatier 1354; holo: P, iso [fragment]: W, LE. 

Illustration: Wu Kuofang, Fl. R. P. Sinicae 13(3): 183, tab. 42 (1997). 

Perennials, (15–) 30–50 (–60) cm tall; rhizome horizontal, creeping, ±short-noded, c. 2 mm 

thick; stem rigid, erect, c. 1.5–2.5 mm thick. Cataphyll usually 0–1; sub-basal leaves 1–2. 

Cauline leaves 2–3, unitubular, perfectly septate, usually long (to c. 20 cm), often reaching 

inflorescence; auricles subobtuse, oblong, scarious, usually 1–2.5 mm long. Lower bract 

leaf-like, usually shorter than the inflorescence, rarely to 5 cm and then often exceeding 

inflorescence. Inflorescence rich, often dense to subdense; main branches 3–8, erect to 

suberect; secondary or all branches often ±abbreviated, of many ±obconical, usually 3–10flowered 

heads. Tepals ±equal or inner ones slightly longer, lanceolate, usually distinctly 

3-veined, greenish, later stramineous-brown, acute to mucronate, c. 2.2–3.0 mm long; 

margins narrow, membranous. Stamens (3–) 6; anthers 0.3–0.5 mm long; filaments 0.9–1.4 mm 

long; style 0.2–0.3 mm long; stigmas c. 1.0–1.5 mm long. Capsule oblong-ovoid, ±acutely 

trigonous, unilocular, c. 2.7–3.1 mm long, subabruptly narrowing into a 0.2–0.3 mm rostrum, 

castaneous-brown to pale brown, usually slightly (rarely distinctly) exceeding perianth. 

Seeds ovoid-ellipsoid, apiculate, c. 0.5 × 0.2–0.3 mm, pale brown, longitudinally 

c. 20-striate, finely reticulate; appendages absent. 

Confined to the Far East, from Manchuria and Russian Ussuri region to S Kuril Is., Japan 

and Korea. 31: KUR, PRM. 36: CHM, CHN-SD. 38: JAP, KOR. Wet meadows, pond shores, 

along tracks and streams. Map 372. 

36. CHINA NORTH-CENTRAL: Shandong, Tai Ping Kung, Lao Shan, C.Y.Chiao 2823 (K). 38. JAPAN: 

Hokkaido, Kushiro, U.Faurie 8644 (E); Hokkaido, Hakodate, 18 Sep 1887, U.Faurie 3209 (K); Hokkaido, 

Hakodate, 29 Aug 1902, U.Faurie 5235 (E); Otaru, 19 Sep 1886, U.Faurie 1365 (K); Hokkaido, Morozan, Sep 

1904, U.Faurie 5727 (W); Honshu, Katakai in Kadzusa, 10 Sep 1953, I.Ito & T.Koyama 973 (W); Honshu, 

Kadzusa, Katakai, I.Ito & T.Koyama TSM973 (K, NSW, TNS); Tokyo, Aug 1905, T.Terasaki (K). 

227

228 


180. Juncus papillosus Franch. & Sav., Enum. Pl. Jap. 2: 98 & 532 (1876) 

T: [Japan] Nippon, in montibus Hakone, P.A.L.Savatier 1355; syn: P; [Japan] circa Yokoska, P.A.L.Savatier 

2521; syn: P. 

Juncus niponensis Buchenau, Bot. Jahrb. Syst. 12: 340 (1890). T: [Japan] Nippon, Yokohama, 

C.J.Maximoxicz; syn: K, LE, PE; [Japan] Tokio, R.Yatabe; syn: n.v. 

Juncus umbellifer H.Lév. & Vaniot, in H.Léveillé, Bull. Soc. Bot. France 51: 292 (1904). T: Corea, Ouen- 

San, Aug 1901, U.Faurie 895; holo: E; iso: P. 

Juncus niponensis var. hakodatensis H.Lév., Feddes Repert. Spec. Nov. Regni Veg. 10: 352 (1912). 

T: Japonia, Yezo [Hokkaido], circa Hakodate, 29 Sep 1902, U.Faurie 5236; syn: E, n.v., P, n.v. 

Juncus nikkoensis Satake, J. Fac. Sci. Univ. Tokyo, Sect. 3, Bot. 4: 185, fig. 20 (1933). T: [Japan] Honshu, 

Jumoto, Nikko, prov. Simotuke, Aug 1931, J.Momiyama; lecto: TI, designated here [the original type sheet 

bears a mixture of J. papillosus and J. wallichianus; left hand plant marked A is the lectotype]. 

Juncus nikkoensis var. minor Satake, J. Fac. Sci. Univ. Tokyo, Sect. 3, Bot. 4: 185 (1933). T: [Japan] 

Honshu, Mt Gandoyama, Prov. Rikuzen, 6 Sep 1931, S.Murai; holo: TI. 

Illustrations: V.I.Krechetovich & N.F.Goncharov, in V.L.Komarov, Fl. SSSR 3: 548, tab. 30 (1935); 

V.S.Novikov, in S.S.Kharkevich, Sosud. Rast. Soviet. Dal. Vost. 1: 71, tab. 5 (1985); Wu Kuofang, Fl. R. P. 

Sinicae 13(3): 187, tab. 43 (1997). 

Map: V.S.Novikov, in S.S.Kharkevich, Sosud. Rast. Soviet. Dal. Vost. 1: 77, fig. 22 (1985) [Russian Far East]. 

Perennials, c. 20–40 cm tall, caespitose to loosely caespitose; rhizome short, ascending; 

stolons short, ascending. Cataphylls 0–2, often with a developed blade, reddish brown. 

Cauline leaves 2–4, terete or slightly compressed, unitubular, with ±obtuse apex, perfectly 

septate, to 12 cm long, 1–2 mm wide; auricles ±obtuse, scarious, c. 0.4–1.0 mm long. Lower 

bract reduced, c. 1–2 cm long, leaf-like, much shorter than inflorescence. Inflorescence rich, 

decompound to supradecompound, loose and diffuse, less often subdense; main branches 

suberect, of many (1–) 2–3 (–5)-flowered, sometimes subcontracted heads. Tepals unequal; 

inner tepals conspicuously longer, ±lanceolate, c. 2.0–2.4 mm long, acute to acuminate, with 

distinct membranous margins; outer tepals to 2.0 mm long, linear-lanceolate, greenish, often 

suffused pale purplish, later ±stramineous-brown. Stamens 3, shorter than or equalling 

tepals; anthers c. 0.4–0.5 mm long; filaments 0.8–1.3 mm long; style c. 0.3–0.5 mm long, 

difficult to distinguish from rostrum apex; stigmas c. 0.6–1.0 mm long. Capsule unilocular, 

narrowly oblong-ovoid, acutely trigonous, c. 3.2–4.0 mm long, acuminate, with a c. 0.3–0.5 mm 

rostrum, thin-walled, pale brownish stramineous, ±shiny, much exceeding perianth. Seeds 

ellipsoid, apiculate, c. 0.5 × 0.2 mm, pale brown, longitudinally c. 16–20-striate, 

transversally reticulate; appendages absent. 

Distributed from E China to the Russian Far East and Japan. 31: AMU, KHA, KUR, PRM, 

SAK. 36: CHI, CHM, CHN, CHS-HE. 38: JAP, KOR. Wet meadows, sandy and gravelly 

shores, along paths. Map 373. 

31. PRIMORYE: Nikolsk-Ussuriysk, 12 Jul 1920, Á.Kiss (PR); S Ussuri region, Pashkowa, V.L.Komarov 354 

(K, LE, P). 36. MANCHURIA: Kirin [Jilin], O-mu Hsien, H.W.Kung 2188 (S); Kirin, between Hui Fa R. 

and Kirin, 1886, H.E.M.James (K). CHINA NORTH-CENTRAL: Shandong, Chefoo Mtns, 1889, E.Faber 

(K). 38. JAPAN: Honshu, Hyogo, Tonomine, Kanzaki-gun, 800 m, K.Iwatsuki & H.Koyama [Pl. Jap. Exs.] 

798 (KYO, LD); Honshu, Aomori, Oipe, Shimokita, H.Koyama & N.Fukuoka [Fl. Jap. Exs.] 223 (BM); 

Honshu, Musashi, Nishitama-gun, Shiofune, M.Mizushima 15546 (S); Honshu, Gunma Pref., Mt Arafune, Oct 

1954, I.Kato (TI); Honshu, Mt Fuji, 21 Aug 1932, K.Hisauti (TI); Hokkaido, Otaru, U.Faurie 185 (BM). 

KOREA: Ouen-san, U.Faurie 895 (P); Quelpaert, U.Faurie 2250 (P). 

181. Juncus virens Buchenau, in H.G.A.Engler, Pflanzenr. (iv.36) 25: 220 (1906) 

Juncus papillosus var. virens (Buchenau) Vorosch. [Fl. Soviet. Dal. Vost. 112 (1966), nom. inval.]; in 

A.K.Skvortsov, Florist. Issled. Raz. Raion. SSSR 157 (1985). T: [Russian Far East] Blagowjestschensk im 

Amurgebiete, 7 Aug 1898, F.K.Karo 89; lecto: LE, fide V.N.Novikov, Nov. Sist. Vyssh. Rast. 19: 60 (1982); 

iso: G, K, L, PR, W [The W specimen probably should be given preference because a picture and a short 

description was provided by Buchenau]. 

Illustration: V.I.Krechetovich & N.F.Goncharov, in V.L.Komarov, Fl. SSSR 3: 548, tab. 30 (1935). 

Map: V.S.Novikov, in S.S.Kharkevich, Sosud. Rast. Soviet. Dal. Vost. 1: 78, fig. 23 (1985) [Russian Far East].


Perennials, 10–40 cm tall, caespitose; rhizome short, ±creeping; stem terete, erect to 

ascending, c. 1.0–1.5 mm in diam. Cataphylls 0–2, short, brownish-greenish. Cauline leaves 

2–3, c. 4–12 cm long, thin, c. 0.6–0.9 mm wide, unitubular, perfectly septate; auricles obtuse 

to truncate, c. 0.4–0.8 mm long, ±scarious. Lower bract leaf-like, c. 2–4 cm long, shorter 

than inflorescence. Inflorescence decompound, ±diffuse; main branches 1–3, erect to erectopatent, 

of 8–40 small usually obconical to semiglobose, (1–) 2–4 (–5)-flowered heads. Tepals 

±unequal, greenish or suffused reddish, with broad scarious margins; outer tepals lanceolate, 

3-veined, c. 1.8–2.0 mm long, acute to mucronate; inner tepals longer, broader lanceolate, to 

2.3 mm long, obtuse. Stamens 3; anthers c. 0.4–0.5 mm long; filaments c. 0.8–0.9 mm long; 

style 0.1–0.2 mm long; stigmas c. 1.0–1.3 mm long. Capsule unilocular, trigonous-ovoid, 

c. 2.5–2.9 mm long, acuminate, with a c. 0.2–0.4 mm rostrum, pale brownish to stramineous, 

±dull, much exceeding perianth. Seeds ellipsoid, apiculate, c. 0.5 × 0.2 mm, pale brown, 

longitudinally c. 16–20-striate, transversally reticulate; appendages absent. 

Confined to the Far East, from the Amur region to E China and N Korea and Japan. 31: AMU, 

KHA, PRM. 36: CHM. 38: JAP, KOR. Meadows along streams, shores. The Japanese 

specimen cited is not quite typical; additional material is needed. Map 374. 

31. AMUR: Zea, Jul 1900, F.K.Karo s.n. (LE, NSW). 36. MANCHURIA: NE Manchuria, Shichotez railway 

station, 1 Aug 1902, D.Litvinov (LE). 38. JAPAN: Honshu, Iwate, Isawa-gun, Mt Yakeishi-dake, 

1100–1400 m, 24 Jul 1958, M.Furuse (S). 

182. Juncus heterophyllus Dufour, Ann. Sci. Nat. (Paris) 5: 88 (1825) 

Juncus lampocarpus var. heterophyllus (Dufour) Duby, Bot. Gall. 1: 477 (1828). T: France, Saint-Sever, 

L.Dufour 71; syn: G-DC; other possible authentic material: Bordeaux, L.Dufour (P). 

Juncus atlanticus Laharpe, Essai Monogr. Jonc. 40 (1825). T: [Algeria] Barbarie [collector not given]; 

holo: G-DEL, n.v. 

Illustrations: F.Buchenau, in H.G.A.Engler, Pflanzenr. (iv.36) 25: 174, fig. 87 (1906); R.Maire, Fl. Afr. Nord 

4: 288, fig. 718 (1957); S.Pignati, Fl. Ital. 3: 439 (1982); M.C.Fernández-Carvajal, in B.Valdés et al. (eds), 

Fl. Vasc. Andalucia Occid. 3: 223 (1987). 

Perennials, usually submersed aquatic, (10–) 20–100 cm long or tall, without rhizome, with 

fasciculate roots sometimes having tubercles. Stems floating or creeping and rooting at 

nodes, branched. Submerged leaves long, narrow, filiform, often overwintering; emergent 

leaves 10–24 cm long, 2–6 mm in diam., unitubular, visibly perfectly septate and nodose; 

sheaths lax; auricles large, obtuse, 1–3.5 mm long, to 2.5 mm wide, ±membranous. Lower 

bract leaf-like, usually shorter (less often equalling or longer) than inflorescence. 

Inflorescence anthelate, 2.5–10 cm long, lax, of (2–) 4–15 (–20) heads each with 2–6 

flowers; pedicels c. 1.0–1.5 mm long; peduncles often flexuous. Tepals 3.5–6 mm long, 

equal or the inner slightly longer than the outer, lanceolate-ovate, obtuse to subobtuse, 

castaneous-coloured or brown-reddish, with broad scarious (later inrolled) margins. Stamens 

6, more than 1/2 as long as tepals; anthers 1.5–2.5 mm long, 2–4 times as long as the 

filaments c. 0.6–0.7 mm long; style 0.5–0.8 (–1.3) mm long; stigmas 1.5–2.0 (–3.0) mm 

long, reddish. Capsule 5–7 mm long, exceeding perianth, trigono-ovoid, subacute, with a 

1–1.5 mm rostrum, unilocular, paler castaneous-coloured or brown-reddish, shiny. Seeds 

±ovoid, 0.5–0.6 (–0.7) × 0.3–0.4 mm, striate, reticulate. 

SW Europe; NW Africa. 12: COR, FRA, POR, SAR, SPA. 13: ITA, SIC. 20: ALG, MOR, 

TUN. In temporarily exposed pool bottoms or lake shores and other periodically inundated 

muddy or sandy basins, on siliceous substrates. Map 375. 

12. FRANCE: Gironde, La Teste-de-Buch, E.J.Neyraut [I.Dörfler, Herb. Norm.] 3869 (PR, W, WU). 

PORTUGAL: Aguiar da Beira, Poço Negro, Jul 1890, M.Ferreira (COI); Tras-os-Montes, Barroso, Lameiro 

Grande, 950 m., 23 Jun 1943, G.Pedro & M.Myre (MA19278). SARDEGNA: Cagliari, between Pula and 

Domus de Maria, H.Glück & H.Poeverlein [A.Kneucker, Cyper. Junc. Exs.] 261 (PR, PRA). SPAIN: 

Badajoz, Herrera del Duque, Valle del Rosal, 21 Jun 1969, S.Rivas Goday & M.Ladero (FCO6473, 

MAF88025); Huelva, entre Tharsis y Villanueva de las Cruces, 13 Apr 1960, S.Rivas Goday, E.F.Galiano & 

S.Rivas Martínez (MA175381, SEV5930). 13. ITALY: Pisa, 1840, P.Savi (PR). 20. ALGERIA: 

Constantine, Mt Ouach, 23 Jun 1934, R.Maire (P). MOROCCO: Atlas Rifain, Isaguen, E.M.Sennen & 

Mauricio [E.M.Sennen, Pl. Esp.] 1580 (W); R.Maire (P); Oulmés, Zaian Mtns, G.Samuelsson 7400 (S). 

229

230 


183. Juncus emmanuelis A.Fern. & J.G.García, Bol. Soc. Brot. ser. 2, 21: 6 (1947) 

T: Portugal, Vale do Falagueiro prope Vendas Novas in Transtagana, Jun 1946, J.Matos; holo: COI. 

Juncus emmanuelis var. inflatifolius A.Fern. & J.G.García, Mem. Soc. Brot. 4: 32 (1948). T: [Portugal] Vale 

do Falagueiro pr. Vendas Novas in Transtagana, Jun 1946, J.Matos ; holo: COI. 

Illustrations: A.Fernandes, J.Garcia & R.Fernandes, Mem. Soc. Brot. 4: tab. I (1948); M.C.Fernandez-Carvajal, 

Anales Inst. Bot. Cavanilles 34: 127–128, fig. 1–3 (1977); M.C.Fernández-Carvajal, in B.Valdés et al. (eds), 

Fl. Vasc. Andalucia Occid. 3: 224 (1987). 

Map: M.C.Fernandez-Carvajal, Anales Inst. Bot. Cavanilles 34: 130 (1977). 

Perennials, (21–) 25–50 (–60) cm tall, with filiform roots producing tuberous thickenings 

10–15 × 3–5 mm; rhizome densely branched, with tufted stems, tufts connected with long 

stoloniform rhizome shoots with internodes 1–5 cm. Stem erect, 2–3 mm in diam, tuberous at 

base, with 2–3 basal sheaths and 2–4 (–5) cauline leaves. Leaves usually 5–20 cm long, 

(1.5–) 2–4.5 (–6) mm wide, unitubular, perfectly septate (septa distinct externally); auricles 

c. 2 mm long, oblong, obtuse, scarious. Inflorescence with 1–5 main branches, often with 1–3 

secondary branches, of (2–) 3–8 (–12) globose or semiglobose heads, each with (3–) 5–30 (–50) 

flowers. Lower bract usually 1–3 cm long, leaf-like, less than 50% inflorescence length. 

Tepals (4–) 4.5–6 mm, equal or the inner ones slightly shorter, narrowly lanceolate or 

subulate, acuminate, often recurved apically, usually ±reddish, with narrow scarious margin; 

central band with (1–) 3–5 prominent veins. Stamens 6, 1/2– 2/3 as long as tepals; anthers 

1.5–2.9 mm long, 4–6 times as long as 0.3–0.5 mm filaments; style c. 1.2 mm long; stigmas 

exserted, 1–2 mm long. Capsule unilocular, 4.0–5.5 mm long, equalling or shorter than 

perianth, narrowly trigonous-ovoid, with a c. 1.0–1.5 mm rostrum, stramineous to brownish, 

shiny. Seeds ovoid, apiculate, (0.5–) 0.6–0.7 × c. 0.3 mm, reticulate, castaneous or 

ferrugineus. 2n = 40, fide A.Fernandes & J.G.García, Bol. Soc. Brot. ser. 2, 21: 7 (1947). 

SW Europe (C, S Portugal, SW Spain). 12: POR, SPA. Margins of watercourses and pools of 

oligo-mesotrophic waters. Map 376. 

12. PORTUGAL: Montargil, Jun 1883, J.Cortezão (COI); Setubal, estrada d’Algeruz, Jun 1901, A.Luisier 

(COI); Vendas Novas, Barranco da Malhada das Vacas, 12 May 1947, A.Fernandes & F.Sousa (COI); 10 Jun 

1946, J.Matos 2307 (COI, S); Vendas Novas, Vale do Falagueiro 10 Jun 1946, J.Matos (COI); Vendas Novas, 

Linha Velha, pr. Adegas, 12 May 1947, A.Fernandes & F.Sousa (COI). SPAIN: Huelva, Almonte, Reserva 

Biológica de Doñana, El Martinazo, 13 May 1966, E.F.Galiano & I.Novo (SEV27506); Huelva, Reserva 

Biológica de Doñana, cerca del Pinar del Raposo, 2 Jun 1977, M.Costa & E.Valdés (FCO6465, SEV59875); 

Huelva, Mazagón, Laguna de las Pozas, 17 May 1979, S.Silvestre, S.Talavera & al. (SEV). 

184. Juncus acutiflorus Ehrh. ex Hoffm., Deutschl. Fl. 125 (1791) 

Juncus foliosus Hoppe, Neues Bot. Taschenb. 1810: 159 (1810), nom. illeg.; Juncus brevirostris Nees, in 

M.J.Bluff & C.A.Fingerhuth, Comp. Fl. German. 1: 435 (1825), nom. illeg.; Juncus acutiflorus var. 

brevirostris Bluff, Nees & Schauer, Comp. Fl. German., ed. 2, 1: 565 (1836); Juncus lampocarpus var. 

acutiflorus (Hoffm.) Neilr., Fl. Wien 97 (1846); Phylloschoenus acutiflorus (Ehrh. ex Hoffm.) Fourr., Ann. 

Soc. Linn. Lyon n. ser. 17: 172 (1869), nom. inval.; Juncus acutiflorus var. genuinus Cout., Bol. Soc. Brot. 8: 

116 (1890), nom. inval.; Juncus acutiflorus subsp. genuinus (Cout.) Cout., Fl. Portugal 118 (1913), nom. 

inval. [synonyms of J. acutiflorus subsp. acutiflorus]; Juncus sylvaticus race brevirostris [Nees] Rouy, Fl. 

France 13: 242 (1912). T: [Germany] Hannoverae, F.Ehrhart [Calam. Tripet. Exs.] 66; lecto: BM, fide 

S.Snogerup, in K.H.Rechinger, Fl. Iranica 75: 24 (1971); iso: LINN-SMITH. 

Juncus acutiflorus var. microcephalus Husn., Joncées 14 (1908). T: not designated. 

Perennials, (20–) 25–110 cm tall, rhizomatous, forming large loose stands. Rhizome 

creeping, stout, sparsely branched, with internodes 0.5–1.5 mm long. Stem erect; cataphylls 

2–3, with uppermost to 20 cm long. Cauline leaves 2–4 (–6), 5–50 cm long, 0.7–2 mm wide, 

unitubular, perfectly septate (septa visible externally, ±distinct); auricles ±firm, scarious, 

small, c. 1.0–1.5 mm long, obtuse. Inflorescence lax or ±condensed, with (8–) 30–80 (–250) 

heads, each of (3–) 5–10 (–20) flowers. Lower bract leaf-like, shorter than inflorescence 

(rarely equal or longer). Tepals unequal; inner tepals (1.8–) 2–2.8 mm long; outer tepals 

shorter, (1.5–) 1.7–2.3 (–2.5) mm long, ±narrowly ovate, apiculate to cuspidate, usually 

slightly recurved at apex, castaneous-coloured at least distally, greenish or pale brown at 

base, sometimes greenish when young, with narrow scarious margin. Stamens 6; anthers 

(0.6–) 0.7–1 mm long, longer than 0.3–0.55 (–0.7) mm filaments; style 0.5–1 mm long;


stigmas 1–1.5 mm long. Capsule (2.3–) 2.5–3.5 (–4) mm long, exceeding perianth, 

unilocular, trigono-ovoid to narrowly pyramidal, gradually narrowed in a 0.5–1 mm rostrum, 

brownish, shiny. Seeds ±ovoid to ellipsoid, 0.4–0.5 (–0.6) × c. 0.2–0.3 mm, c. 25-striate, 

reticulate; appendages absent. 

Europe; NW Africa; SW Asia; Newfoundland; introduced in New Zealand. Two subspecies 

are recognized. 

Stems, basal sheaths and leaves smooth or slightly longitudinally striate 

(mainly in dried specimens) 184a. subsp. acutiflorus 

Stems, basal sheaths and leaves strongly transversally rugose 184b. subsp. rugosus 

184a. Juncus acutiflorus Ehrh. ex Hoffm. subsp. acutiflorus 

Juncus nemorosus Sibth., Fl. Oxon.: 114 (1794), nom. illeg., non Pollich (1776). T: several references to 

possible syntypes; e.g.: Juncus articulatus var. sylvaticus L., Syst. Veg., ed. 12, 250 (1767): Juncus folio 

articulato sylvaticum, V.A.Haller, later referred to as Haller 1323 [V.A.Haller, Hist. Stirp. Helv. 2: no. 1323 

(1768)]; syn: P-Haller; R.Morison, Pl. Hist. Univ. 8, tab. 9, fig. 2 ad dextram [R.Morison, Pl. Hist. Univ. 3, 

sect. 8, tab. 9, fig. 2 (1715); syn [the figure represents a rhizomatous plant very probably belonging to 

J. acutiflorus; one of several stems infested by Livia juncorum]. 

Juncus spadiceus Schreb., in A.F.Schweigger & F.Körte, Fl. Erlang. 149 (1811). T: [Germany] Erlangen, 

J.C.D.Schreber; syn: L. 

Juncus acutiflorus var. minor J.Gay ex Laharpe, Essai Monogr. Jonc. 40 (1825). T: [France] environs du 

Mans et de Montpellier, J.Gay; syn: n.v. 

Juncus acutiflorus var. repens J.Gay ex Laharpe, Essai Monogr. Jonc. 40 (1825). T: [locality not indicated], 

J.Gay; syn: n.v. 

Juncus acutiflorus var. pauciflorus Spenn., Fl. Friburg. 1: 184 (1825). T: not designated. 

Juncus acutiflorus var. multiflorus Weihe ex Spenn., Fl. Friburg. 1: 184 (1825). T: A.Weihe [Deutsche 

Gräser] 92; syn: PRC. 

Juncus acutiflorus var. laxus Spenn., Fl. Friburg. 1: 184 (1825). T: not designated. 

Juncus acutiflorus var. viviparus Spenn., Fl. Friburg. 1: 184 (1825). T: not designated [infested by Livia 

juncorum Latreille]. 

Juncus acutiflorus β [unranked] hagenbachianus Gaudin, Fl. Helv. 2: 551 (1828). T: [Switzerland] 

Basileae, C.F.Hagenbach; holo: n.v. 

Juncus acutiflorus γ [unranked] tenuifolius Gaudin, Fl. Helv. 2: 551 (1828). T: [Switzerland] prope 

Promenthou, J.P.Monnard; holo: n.v. 

Juncus acutiflorus var. exoletus Rchb., in J.C.Mössler & H.G.L.Reichenbach, Handb. Gewächsk., ed. 3, 1: 607 

(1833). T: not designated. 

Juncus acutiflorus [Spielart] effusus G.Mey., Chloris Han. 566 (1836). T: icon in T.Host, Icon. Descr. 

Gram. Austriac. 3: tab. 86 (1805); syn; Göttingen, Bollensen, G.Meyer; syn: n.v. 

Juncus sylvaticus var. macrocephalus W.D.J.Koch, Syn. Fl. Germ. Helv. 729 (1837); Juncus sylvaticus var. 

major Parl., Fl. Ital. 2: 337 (1852), nom. illeg. T: not designated. 

Juncus sylvaticus var. pallescens W.D.J.Koch, Syn. Fl. Germ. Helv. 729 (1837); Juncus sylvaticus subvar. 

pallescens (W.D.J.Koch) Rouy, Fl. Fr. 13: 242 (1912). T: not designated. 

Juncus sylvaticus [unranked] virescens Döll, Fl. Baden 1: 329 (1857). T: Baden, Oberbruch, J.C.Döll; syn: n.v. 

Juncus sylvaticus var. congestus Lange, Vidensk. Meddel. Naturhist. Foren. Kjøbenhavn 2: 67 (1860). 

T: [Spain] ad Santiago, 29 Aug 1851, J.M.C.Lange; holo: C, n.v. [misspelled as confertus in M.Willkomm & 

J.M.C.Lange, Prodr. Fl. Hisp. 1: 185 (1861)]. 

Juncus sylvaticus var. viviparus Lange [Vid. Medd. Naturk. Foren. Kjφbenhavn: 67 (1860), nom. inval.] ex Lange, in 

M.Willkomm & J.Lange, Prodr. Fl. Hisp. 1: 185 (1861). T: Santander, Oct 1852, J.Lange; syn: COI. 

Juncus rufus Miel. ex Huebener, Flora 22: 490 (1839), nom. inval. [not accepted by the author]; Juncus 

acutiflorus II. [unranked] rufus Asch. & Graebn., Syn. Mitteleur. Fl. 2 (2): 468 (1904). T: [Austria] 

Zellersee in Pinzgau, M.Mielichhofer, n.v. 

Juncus acutiflorus B. [unranked] multiflorus Asch. & Graebn., Syn. Mitteleur. Fl. 2 (2): 468 (1904). 

T: not designated. 

Juncus acutiflorus III. [unranked] wulfenii Asch. & Graebn., Syn. Mitteleur. Fl. 2 (2): 468 (1904). T: not 

designated. 

231

232 


Juncus sylvaticus var. paniculatus Lojac., Fl. Sicul. 3: 158 (1909). T: [Sicily, otherwise not indicated]. 

Juncus sylvaticus var. virescens Lojac., Fl. Sicul. 3: 158 (1909), nom. illeg. T: [Sicily, otherwise not 

indicated] A.Todaro; syn: n.v. 

Juncus sylvaticus var. brachiatus Lojac., Fl. Sicul. 3: 158 (1909). T: [Sicily, otherwise not indicated]. 

Juncus acutiflorus [Spielart] coarctatus G.Mey., Chloris Han. 566 (1836), nom. inval. 

[Juncus sylvaticus auct., non Reichard (1778): F.G.T.Rostkovius, De Junco 40 (1801)] 

Illustrations: T.Szynal & J.Mądalski, Atlas Fl. Pol. 2(2): 143 (1931); Ö.Nilsson & S.Snogerup, Bot. Not. 125: 

132, fig. 69 (1972); S.Ross-Craig, Drawings Brit. Pl. 8 (30): tab. 17 (1973); M.C.Fernández-Carvajal, in 

B.Valdés et al. (eds), Fl. Vasc. Andalucia Occid. 3: 225 (1987). 

Maps: E.Hultén, Kongl. Svenska Vetenskapsakad. Handl., ser. 5, 7(1): map 138 (1958); H.Meusel, E.Jäger & 

E.Weinert, Vergl. Chorol. Zentraleur. Fl. 1: 85 (1965); E.Hultén & M.Fries, Atlas N.Eur. Vasc. Pl. 1: 94, map 

187 (1986). 

Plants with stems, basal sheaths and leaves smooth or slightly longitudinally striate (mainly 

in dried specimens). No ridges in outline of transverse section of stems and leaves, with all 

epidermal cells (1–) 1.5–2 times as long as wide. 2n = 40, fide S.Snogerup, in Ö.Nilsson & 

S.Snogerup, Bot. Not. 125: 131 (1972); M.Zandee, Proc. Kon. Ned. Akad. Wetensch., C, 84: 

250 (1981). Fig. 105. 

Widely distributed in C, W and S Europe, extending to W Ukraine; also NW Africa and E 

Mediterranean, reaching SW Asia; probably introduced in Newfoundland and surely so in 

New Zealand and New South Wales. Reported from Turkey and Iran but no material seen 

and the occurrence remains doubtful. 10: DEN, fin (extinct), FOR, GRB, IRE, NOR, swe 

(extinct). 11: AUT, BGM, CZE, GER, NET, POL, SWI. 12: BAL, COR, FRA, POR, SAR, 

SPA. 13: ALB, BUL, GRC, ITA, ROM, SIC, YUG. 14: kry, UKR. 20: MOR. 34: IRQ, LBS. 

50: nsw. 51: nzs. 72: lab, nfl. Meadows on oligotrophic to moderately nutrient-rich, alluvial 

or fertilized, wet or damp soils, often inundated at least in winter, and usually mowed or 

grazed. Map 377 (secondary distribution not shown). 

10. GREAT BRITAIN: East Norfolk, between Winterton and Winterton Ness, P.D.Sell [Pl. Brit. Exs.] 103 

(K, UPS). 11. CZECHOSLOVAKIA: Šumava Mtns, Horní Vltavice, B.Křísa (LD, PRC). GERMANY: 

Bavaria, Bamberg, Fischer [I.Dörfler, Herb. Norm.] 4782 (PRC, UPS). 12. FRANCE: Charenté, NE of 

Leigne, Villefagnan, E.Contré [Soc. Fr. Éch. Pl. Vasc.) 4711 (P). PORTUGAL: Alto Douro, Murça, serra do 

Ratiço, Jul 1880, M.Ferreira (COI); Tras-os-Montes, Serra do Brunheiro, Chaves, Jul 1892, A.Moller (COI). 

SPAIN: Asturias, Salas, La Espina, 4 Jul 1975, M.Mayor, F.Navarro, T.E.Díaz & M.C.Fernández-Carvajal 

(FCO15693); Pontevedra, Vilaboa, San Adrián, 18 Jul 1970, S.Castroviejo (MA197464). 13. ITALY: 

Calabria, E of S. Giovanni in Fiore, 25 Jun 1980, B.Snogerup & S.Snogerup (LD). 

184b. Juncus acutiflorus subsp. rugosus (Steud.) Cout., Fl. Port. 118 (1913) 

Juncus rugosus Steud., Syn. Pl. Glumac. 2: 298 (1855); Juncus sylvaticus subsp. rugosus (Steud.) Nyman, 

Consp. Fl. Eur. 747 (1882); Juncus acutiflorus var. rugosus (Steud.) Cout., Bol. Soc. Brot. 8: 116 (1890). 

T: [Portugal] Cintra Lusitaniae, C.F.Hochstetter [Herb. It. Lusit.] 331; holo: P. 

Illustrations: M.C.Fernández-Carvajal, Anales Jard. Bot. Madrid 39: 334, fig. 36, 336, fig. 39, 337, fig. 41, 43 

[all anatomy] (1983); M.C.Fernández-Carvajal, in B.Valdés et al. (eds), Fl. Vasc. Andalucia Occid. 3: 225 

(1987). 

Map: M.C.Fernández-Carvajal, Anales Jard. Bot. Madrid 39: 339, fig. 44 (1983). 

Plants with stems, basal sheaths and leaves strongly transversely rugose. Transverse section 

of stems and leaves with ridges in outline, having some epidermal cells (1–) 1.5–2 times as 

long as wide, alternating with other larger ones 3–5 times as long as wide. 

Confined to SW Europe (SW Spain; C & S Portugal). 12: POR, SPA. Meadows and heaths 

developed on oligotrophic to moderately nutrient-rich, alluvial or fertilized, wet or damp 

soils, often inundated at least in winter. Map 378. 

12. PORTUGAL: Algarve, Pico de Foia, 11 Jun 1960, A.Fernandes, R.Fernandes & J.Matos (COI); Alto 

Alentejo, Vendas Novas, Barranco da Malhada das Vacas, 12 May 1947, A.Fernandes & F.Sousa (COI); Beira 

Litoral, Pinhal do Urso, Jul 1898, M.Ferreira (COI, MA19326); Estremadura, entre Aguas de Moura y 

Setubal, 30 m, 18 May 1990, T.E.Díaz (FCO17870). SPAIN: Huelva, Doñana, Caño de la Raya, 18 Jun


Figure 105. Juncus acutiflorus. A, habit; B, flower head; C, perianth with capsule; 

D, capsule T.S.; E, capsule; F, outer tepal; G, inner tepal; H, auricles; I, stem T.S.; J, leaf blade 

T.S. (A–J, P.F.F.Schulz, Aug 1901, KRA). Scale bars: A = 5 cm; B = 2 mm; C–G, I, J = 1 mm; 

H = 3 mm. Drawn by T.Szynal. Reproduced with permission, from T.Szynal & J.Mądalski, 

Atlas Fl. Polsk. 2/2: 143 (1931) 

233

234 


1978, E.Valdés-Bermejo & S.Castroviejo (FCO10389); Huelva, Doñana, Fuente del Duque, 15 Jun 1973, 

B.Cabezudo (SEV17798); Cádiz, Tarifa, Sierra de Saladavieja, El Pedregoso, 200 m, 22 Jul 1980, J.Arroyo & 

Gil (SEV78181). 

185. Juncus valvatus Link, in H.A.Schrader, J. Bot. 1799(2): 316 (1800) 

T: [Portugal] Montachique, J.H.F.Link; holo: B, probably destroyed. 

Juncus echinuloides Brot., Fl. Lusit. 1: 518 (1804); Juncus valvatus var. echinuloides (Brot.) Coss. & Durieu, 

in J.B.G.M.Bory & M.C.Durieu, Expl. Sci. Algér. 2: 269 (1868). T: [Portugal] Lamarosa, et alibi in Beira 

australi, et Extremadura, F.Brotero; syn: LISU, not extant, probably destroyed. 

Juncus caricinus Durieu, in J.B.G.M.Bory & M.C.Durieu, Expl. Sci. Algérie Fl. 1: tab. 43, fig. 1 (1849); 

Juncus valvatus var. caricinus (Durieu) Coss. & Durieu, in J.B.G.M.Bory & M.C.Durieu, Expl. Sci. Algérie Fl. 

2: 269 (1868). T: [Algeria] Constantine, M.C.Durieu; syn: K [herb. J.Gay], P. 

Juncus valvatus subsp. conimbrigensis A.Fern. & R.B.Fern., Flórula Vasc. Mata Bufarda 154 (1991). 

T: Portugal, Coimbra, Mata, 1 Jun 1988, A.Fernandes, R.B.Fernandes & al. s.n.; holo: COI. 

Illustrations: L.C.Trabut, in J.A.Battandier & L.C.Trabut, Fl. Algérie, ed. 2, 88, tab. 2 (1895); R.Maire, Fl. Afr. 

Nord 4: 291, fig. 720 (1957); M.C.Fernández-Carvajal, Anales Jard. Bot. Madrid 39: 341, fig. 45 (1983). 

Perennials, 11–50 cm tall. Rhizome horizontal creeping, short, short-noded; internodes 

usually 1–3, rarely to 7 mm long. Stem erect, terete, 1–2 mm in diam. Cataphylls 1–3, lax, 

usually 3–6 cm long, stramineous-brown; basal leaves 1–3. Cauline leaves 2–3 (–4); blade 

usually 5–15 cm long, 1–2 mm in diam., terete somewhat laterally compressed, unitubular, 

perfectly septate (septa externally visible, not very distinct); sheaths of cauline leaves usually 

lax and ±dilated; auricles scarious to yellowish, oblong, obtuse or subacute, 1.0–1.5 mm long. 

Lower bract leaf-like, 0.5–3.0 cm long, usually erecto-patent, equalling or longer than 

inflorescence (rarely shorter). Inflorescence of 1–5 (–7) globose, many-flowered congested 

heads, or 1–2 heads on erect to suberect rigid peduncles. Tepals subequal, with outer ones 

(3.5–) 3.7–4.5 mm long, slightly longer than inner ones, very narrowly triangular-lanceolate, 

subulate at apex, greenish dorsally, pale brown at apex,with central band distinctly 3-veined. 

Stamens 3, c.1/2 as long as perianth; anthers 0.5–0.7 (–0.8) mm long, 2 /3– 1 /2 as long as 0.8–1.0 mm 

filaments; style c. 0.2 mm long; stigmas c. 0.7–1.5 mm long. Capsule 4–5 mm long, 

exceeding perianth, unilocular, narrowly pyramidal, gradually tapering to a 0.5–1 mm 

rostrum, yellow brownish, shiny. Seeds (0.3–) 0.4–0.5 mm long, ellipsoid, finely reticulate. 

2n = 40, fide M.Zandee, Proc. Kon. Ned. Akad. Wet., C, 84: 245 (1981). 

SW Europe (C & S Portugal); N Africa (Algeria, Tunisia). 12: POR. 20: ALG, TUN. Wet 

meadows, margins of watercourses and other periodically inundated places. Map 379. 

12. PORTUGAL: Coimbra, Eiras, Jun 1892, M.Ferreira (MA19296); M.Ferreira [Fl. Lusit. Exs.] 53 (WU); 

M.Ferreira [A.Kneucker, Cyper. Junc. Exs.] 82 (PR, PRA, WU); Quinta do Venâncio, arredores de Leiria, 

20 May 1952, A.Fernandes, R.Fernandes & F.Sousa (COI); Miranda do Corvo, Jun 1883, B.F. de Mello (COI); 

Vermoil, Jul 1890, A.Moller; Praia das Maças, 20 May 1963, J.Matos & M.C.Alves (COI, MA188338); Mafra, 

Tapada Real, Sep 1885, O.Simões (COI); entre Thomar e Ferreira do Zêzere, May 1914, M.Ferreira (COI). 

20. ALGERIA: Lambèse, B.Balansa [Pl. Algér.] 737 (PR). 

186. Juncus fontanesii J.Gay ex Laharpe, Essai Monogr. Jonc. 42 (1825) 

Juncus fontanesii var. typicus Buchenau, in H.G.A.Engler, Pflanzenr. (iv.36) 25: 192 (1906), nom. inval. 

T: [Algeria; label text by J.Gay: Ex herbario suo atlantico dedit Desfontaines 21 Jul 1822] Barbarie, 

R.Desfontaines; holo: K; iso: P-DESF. 

Perennials; rhizome absent or weakly developed, short; above-ground stolons often present. 

Stems erect 5–40 cm tall; cataphylls 1–3. Cauline leaves 2–5. Leaves unitubular, perfectly 

and distinctly septate, terete or slightly canaliculate at base; auricles scarious, to 3 mm long. 

Lower bract leaf-like, shorter than inflorescence. Inflorescence of (1–) 3–10 (–15) 

semiglobose to globose heads, each with (4–) 6–20 (–25) flowers. Tepals (2.4–) 3–4 mm 

long, subequal or the inner ones slightly longer than the outer, narrowly ovate, acute to 

acuminate, distinctly 3-veined, green-brownish or reddish, with narrow scarious margin. 

Stamens 6, 1/3– 1/2 as long as perianth; anthers 0.7–1.4 mm long, 1.5–3 times as long as 

filaments. Style 1–1.5 mm long. Capsule (3–) 4–5.8 mm long, exceeding perianth, 

unilocular, ovoid-trigonous or narrowly pyramidal, tapering to 0.4–1.5 (–2.0) mm mucro,


brownish to ferrugineous, shiny. Seeds 0.4–0.55 (–0.6) mm, pyriform to ovoid, shortly 

apiculate, longitudinally 20–25-striate, transversally reticulate; appendages absent. 

S Europe; N Africa; W Asia, extending to Pakistan. A variable species with a series of local 

forms; four subspecies are recognized but intermediates are known. 

1 Plants mat-forming, with short branched ascending rhizomes; tepals 

shorter than 3 mm 186d. subsp. brachyanthus 

1: Plants caespitose or with long above-ground stolons, not mat-forming; 

rhizomes absent or short, sparsely branched, not ascending; tepals 

usually 3 mm or longer 

2 Rhizome absent; stolons very long, to 2 m; capsule distinctly exceeding 

perianth, trigonous-ovoid, tapering into a mucro 1–2 mm long 186a. subsp. fontanesii 

2: Rhizome present, short; stolons short or absent; capsule slightly 

exceeding perianth, narrowly pyramidal to trigonous-ovoid, abruptly or 

±gradually contracted into a mucro 0.4–1.0 (–1.5) mm long 

3 Plants usually 15–40 cm tall; tepals narrowly ovate, with narrow 

scarious margins; capsule with a mucro 0.5–0.9 (–1.5) mm long 186b. subsp. pyramidatus 

3: Plants usually 8–15 (rarely to 25) cm tall; tepals ovate, with broad 

scarious margins; capsule with a mucro 0.4–0.8 mm long 186c. subsp. kotschyi 

186a. Juncus fontanesii J.Gay ex Laharpe subsp. fontanesii 

Juncus repens Req., in J.Guérin, Descr. Font. Vaucluse, ed. 2, 253 (1813), nom. illeg., non Michx. (1803). 

T: [France] Cadenet, 1813, E.Requien; syn: G; [authentic material: Avignon, E.Requien (K)]. 

Juncus lagenarius J.Gay ex Laharpe, Essai Monogr. Jonc. 42 (1825); Juncus striatus var. lagenarius 

(Laharpe) Parl., Fl. Ital. 2: 344 (1852); Juncus striatus subsp. lagenarius (Laharpe) Arcang., Comp. Fl. Ital. 

716 (1882). T: [France; text written by J.Gay:] Robert misit Decembr. 1823. Teloni [Toulon] lectum 1822; 

lecto: K, herb. J.Gay, designated here. 

Juncus striatus var. pauciflorus Tineo ex Parl., Fl. Ital. 2: 344 (1852). T: [Italy] Sicilia, Monti di Madonia, 

V.Tineo; syn: FI, n.v. 

Juncus hochstetteri Steud., Syn Pl. Glumac. 2: 299 (1855). T: [Spain] C.F.Hochstetter [Herb. Un. It. Lusit.] 

334; syn: P, n.v. 

Juncus striatus var. diffusus Lange, in M.Willkomm & J.Lange, Prodr. Fl. Hispan. 1: 184 (1861). 

T: [Italy, Naples] Castel-di-Sangro, Aprutii, 14 Jul 1856, E.Huet du Pavillon & A.Huet du Pavillon [Pl. 

Neapol. Exs.] 432; syn: K. 

Juncus duvalii Loret, in H.Loret & A.Barrandon, Fl. Montpellier 685 & 848 (1875), n.v.; Loret, Rev. Soc. Sci. 

Nat. 4: 56 (1885). T: n.v. 

Juncus fontanesii var. distortus Lojac., Fl. Sicul. 3: 159 (1909). T: [Sicily, otherwise not indicated]. 

Juncus webbianus V.I.Krecz., Izv. Bot. Sada Akad. Nauk S.S.S.R 30: 98 (1932). T: [Spain, Granada] 

Hispania, prope Granatam, 1849, P.E.Boissier & G.F.Reuter; holo: LE. 

Juncus pisuergae Losa & P.Monts., Anales Inst. Bot. Cavanilles 10(2): 458 (1952). T: [Spain] Orillas del 

Pisuerga, por encima de Cervera, sitios húmedos o encharcados, Jul 1949, M.Losa & P.Montserrat; syn: 

BCF4417 

Juncus stellatus Willd. ex Kunth, Enum. Pl. 3: 329 (1841), nom. inval. 

Illustrations: F.Buchenau, in H.G.A.Engler, Pflanzenr. (iv.36) 25: 191, fig. 93 (1906); R.Maire, Fl. Afr. Nord 

4: 294, fig. 722 (1957); S.Pignati, Fl. Ital. 3: 441 (1982). 

Plant with stolons up to 200 cm long. Rhizome usually absent. Stems erect, 5–40 cm tall. 

Cataphylls 1–3; cauline leaves 2–5. Inflorescence with (1–) 3–10 (–15) semiglobose to 

globose heads, each with (4–) 6–20 (–25) flowers. Lower bract to 2 cm long. Tepals (2.4–) 3–4 × 

0.6–0.9 mm, subequal or the inner ones slightly longer, narrowly ovate, acute to acuminate, 

green-brownish or reddish, with narrow scarious margin. Anthers 0.7–1.4 mm long, 1.5–3 

times as long as filaments; style 1–1.5 mm long; stigmas to c. 1 mm long. Capsule (3–) 4–5.8 mm 

long, exceeding perianth, unilocular, ovoid-trigonous, tapering to a 1–1.5 (–2) mm rostrum, 

brownish to ferrugineus, shiny. Seeds 0.4–0.55 (–0.6) mm long, pyriform to ovoid, shortly 

apiculate. 2n = 80, fide M.Zandee, Proc. Kon. Ned. Akad. Wetensch., C, 84: 245 (1981) 

[vouchers not seen]. 

235

236 


S Europe and N Africa, W Turkey. Introduced in Australia. 12: BAL, FRA, POR, SAR, SPA. 

13: ALB, GRC, ITA, SIC, YUG, TUE. 20: ALG, MOR, TUN. 34: TUR. 50: vic. 

Mediterranean humid grasslands of tall grasses and rushes, streamsides and temporarily 

inundated places, mostly 0–1000 m. Map 380 (secondary distribution not shown). 

12. PORTUGAL: Lisboa, Ribeiro de Caparide, Aug 1880, A.X.Pereira Coutinho (COI); Cascaes, Sep 1890, 

A.X.Pereira Coutinho (COI); Vendas Novas, Vale do Falagueiro, 10 Jun 1947, J.Matos (COI). SPAIN: 

Albacete, Nacimiento del Mundo, 6 Sep 1950, S.Rivas Goday & E.Monasterio (SEV5933, MAF29997); 

Teruel, Lagunica de Tornos, 6 Jul 1958, P.Montserrat (FCO6469); Madrid, Puente de los Franceses, Jun 1921, 

A.Aterido (MA146139); Jerez, Dehesa de Malduerme, 10 Jul 1880, J.M.Pérez-Lara (MAF30158). 

13. ALBANIA: Ljushna, 19 Jun 1918, J.Schneider (W). ITALY: Abruzzi, Mt Mozzone, Aug 1906, G.Rigo (W). 

SICILIA: Mt Madonia, Jul 1873, G.Strobl (WU); Palermo, Jun 1906, H.Ross (W). 

Occasional intermediates between subsp. fontanesii and subsp. pyramidatus are found from 

Italy eastwards [e.g., Italy, Calabria, Corregliano, R.Huter, P.Porta & G.Rigo [It. Ital. III] 

598 (W)]. 

186b. Juncus fontanesii subsp. pyramidatus (Laharpe) Snogerup, in K.H.Rechinger, 

Fl. Iranica 75: 25 (1971) 

Juncus pyramidatus Laharpe, Essai Monogr. Jonc. 40 (1825); Juncus fontanesii var. pyramidatus (Laharpe) 

Buchenau, Bot. Jahrb. Syst. 1: 140 (1880). T: Syrie, J.J.H.Labillardière; syn: P; [label text by J.Gay: en 

Egypte par M. Savigny, donnée 1818] Egypt, M.J.C.Savigny; syn: G-DEL, K. 

Juncus camptotropus V.I.Krecz., Izv. Bot. Sada Akad. Nauk S.S.S.R 30: 98 (1932). T: [Turkey] Paphlagonia, 

wilajet Kastambuli, Tossia, ad ripas Devrek-tschai, 17 Jun 1892, P.Sintenis 4248; holo: LE; iso: E, K, LD, PR, W. 

Densely to loosely caespitose; rhizome short, with intravaginal stolons often present and 

above-ground stolons sometimes developed; stem 15–40 cm tall, erect to ascending. 

Cataphylls 1–3 (–4); cauline leaves 3–5, 5–15 cm long, 1.5–2.5 mm wide; auricles 1.5–3 mm 

long, obtuse. Lower bract 0.5–2.5 cm long. Inflorescence usually to 5–7 cm long, of 3–20 

semiglobose to subglobose 15–20-flowered heads; flower pedicels 0.3–1 mm long. Tepals 

±equal, 3–4 × 0.7–0.9 mm, oblong-lanceolate, apiculate to acuminate, green to brownish, often 

suffused red; margins narrow, scarious. Anthers 0.8–1.2 mm long; filaments c. 0.4–0.6 mm long; 

style 0.5–1 mm long; stigmas c. 1.5 mm long. Capsule narrowly pyramidal to subprismatic, 

4.5–5.0 mm long, conspicuously exceeding perianth, gradually narrowing into 0.5–1.2 mm 

rostrum, pale brown to castaneous-brown; seeds 0.4–0.5 × 0.2–0.3 mm, pale brownish. 

E Mediterranean, eastwards to Iraq; N Africa from Tunisia eastwards. 13: ITA, SIC. 20: 

EGY, LBY, TUN. 34: CYP, IRQ, LBS, PAL, SIN, TUR. 35: SAU, YEM. Temporarily wet 

grasslands, along streams, mostly 0–1500 m, in desert areas to 2500 m. Map 381 (isolated 

occurrence in Yemen and S Saudi Arabia not shown). 

20. EGYPT: Faiyum, 5 May 1967, G.Romée (LD); Tamieh near Faiyum, G.A.Schweinfurth 59 (PR). 

TUNISIA: Zakhouan, J.L.Kralik [Pl. Tunet.] 366 (W). 34. CYPRUS: Paphos, Agios Neophytos, 21 Jul 

1939, H.Lindberg (W). IRAQ: N of Shaqlawa, 950 m, Anders 1464 (W). LEBANON: Kartuba, Ard-il- 

Akluk, 650 m, E.Hartmann [A.Kneucker, Cyper. Junc. Exs.] 83 (PR, PRA). PALESTINE: Jordan, 20 km 

S of Ajlun, Jallad, L.Boulos & Lahham (BM); Israel, Wadi Rubin, 10 Jul 1933, N.Feinbrun & J.Amdursky [Fl. 

Palaest. Exs.] 220 (PR). 35. SAUDI ARABIA: between Abha & Jebel Sauda, S.Collenette 2946 (E). 

YEMEN: Yarim, 2600 m, J.R.I.Wood 1814 (BM). 

186c. Juncus fontanesii subsp. kotschyi (Boiss.) Snogerup, in K.H.Rechinger, Fl. 

Iranica 75: 25 (1971) 

Juncus kotschyi Boiss., in K.G.T.Kotschy, Pl. Persiae Austr. [exsiccate series edited by R.F.Hohenacker, 

printed label description], no. 446 (1845) & Boiss., Diagn. Pl. Orient., ser. 1, 7: 101 (1846); Juncus 

pyramidatus var. kotschyi (Boiss.) Boiss., Fl. Orient. 5: 360 (1882); Juncus fontanesii var. kotschyi (Boiss.) 

Buchenau, Bot. Jahrb. Syst. 12: 329 (1890). T: [Iran] m. Sabst-Buschon pr.[ope] u.[rbem] Schiras, 31 May 

1842, K.G.T.Kotschy [Pl. Pers. Austr.] 446; lecto: G-BOISS, fide S.Snogerup, in K.H.Rechinger, Fl. Iranica 

75: 25 (1971); iso: BM, CGE, G, K, P, PR, UPS. 

Juncus arianus V.I.Krecz., Izv. Bot. Sada Akad. Nauk S.S.S.R 30: 99 (1931); Juncus kotschyi var. arianus 

(V.I.Krecz.) Novikov, Novosti. Sist. Vyssh. Rast. 15: 90 (1979). T: [Turkmenistan] Turkestania, Badchyz, 

Tschilduchter, 2 Sep 1930, M.G.Popov; holo: LE.


Caespitose; stem 10–20 (–25) cm tall; rhizome short; stolons mostly intravaginal. Cataphylls 

1–2; cauline leaves 2–4, sometimes exceeding inflorescence, 3–10 cm long, to 2 mm wide; 

auricles 2–3 mm long, scarious. Lower bract to c. 1.5 cm long. Inflorescence of 3–10 (–13) 

usually semiglobose 7–20-flowered heads to c. 1 cm in diam. Tepals ±equal or the inner ones 

slightly longer, 2.9–3.5 mm long, oblong-ovate, greenish to brownish, often suffused 

reddish, ±acute; inner tepals to c. 1.2 mm wide, often mucronate; margins to 0.2 mm wide, 

membranous. Anthers 0.9–1.1 mm long; filaments 0.5–0.6 mm long; style 0.4–0.5 mm long; 

stigma c. 1.0–1.5 mm long. Capsule ovoid-trigonous, c. 3.5–4.0 mm long, exceeding perianth 

or slightly so, subabruptly contracted into 0.3–0.7 mm rostrum, pale brown. Seeds 0.4–0.5 × 

0.2–0.3 mm, pale brown, longitudinally c. 20–24-striate, indistinctly transversally reticulate; 

appendages absent. 

Western Asia, from Lebanon to Afghanistan and Pakistan. 32: TKM. 34: AFG, IRN, IRQ, 

LBS. 40: PAK. Wet meadows near streams, ditches, mostly 1200–2500 m. Map 382. 

34. IRAN: Khorasan, Hazar Masdjid [Masjed], between Ardak and Tolgor [Talqur], 1200–1600 m, 

K.H.Rechinger & F.Rechinger 4910 (W); Khost, Yakubi [Ya'qubi], 1200 m, K.H.Rechinger 35566 (W). 

IRAQ: Kurdistan, Kirkuk, Darband-i Bazian, C.Haussknecht 976 (JE). 40. PAKISTAN: Baluchistan, Hanna, 

2000 m, 15 Jul 1957, Nasin 28415 (K). 

186d. Juncus fontanesii subsp. brachyanthus Trab., in J.A.Battandier & L.C.Trabut, 

Fl. Algérie, ed. 2, 86, tab. 2 (1895) 

Juncus brachyanthus (Trab.) T.Durand & B.D.Jacks., Ind. Kew., Suppl. 1: 230 (1906), nom. inval. 

T: [Algeria, Oran] bords de la Mekerra à Bedeau, L.C.Trabut; syn: n.v. 

Juncus fontanesii var. melanocephalus Trab., in E.Jahandiez & R.Maire, Pl. Marocc. Nov., no. 1 (1923), n.v. 

T: [Morocco], n.v. 

Illustration: L.C.Trabut, in J.A.Battandier & L.C.Trabut, Fl. Algérie, ed. 2, 88, tab. 2 (1895). 

Perennials, mat-forming or stoloniferous; rhizomes branched, thin, short, ascending to 

creeping, subterranean; internodes c. 1–2 cm long; individual tussock formed by stems and 

intravaginal shoots; lower elevation forms with long above-ground stolons. Stem usually 

5–15 cm tall, erect. Basal leaves 1–2, with brownish sheaths, to 5 cm long, c. 0.5–0.8 mm 

wide; cauline leaves 2–3, almost reaching inflorescence, acute, unitubular, perfectly 

distinctly septate. Lower bract reduced, sheath-like with short blade, usually shorter than 

inflorescence, to c. 1 cm long. Inflorescence usually 0.5–3 cm long, of 1–5 (–8) brown to 

blackish castaneous-brown semiglobose 3–10-flowered heads 7–9 mm in diam.; branches 

usually abbreviated; capitulum bracts scarious-whitish, prominently brownish veined. Tepals 

brown or usually blackish castaneous-brown, often with green midrib, ±equal; outer tepals 

carinate, ovate-lanceolate, acuminate, 2.5–3.0 mm long; inner tepals ovate, subacute or 

subobtuse and mucronate, with contrasting whitish-scarious, 0.2–0.3 mm wide margins. 

Stamens 6; anthers c. 1.0–1.4 mm long; filaments 0.3–0.5 mm long; style c. 1.0–1.2 mm 

long; stigmas c. 1.0 mm long. Capsule unilocular, ovoid-trigonous, acute to acuminate, dark 

castaneous-brown, shining, c. 3 mm long, including 0.5–0.8 mm rostrum; rostrum exceeding 

perianth. Seeds ovoid-pyriform, c. 0.45–0.5 × 0.2–0.3 mm, pale brown, apiculate, 

longitudinally 20–24-striate, transversally reticulate; appendages absent. 

Confined to Algeria and Morocco. 20: ALG, MOR. Mountain meadows, wet sites along 

streams, 1500–3000 m. Map 383. 

20. MOROCCO: Moyen Atlas, Tuiouane, 1600 m, 24 May 1924, E.Jahandiez 462 (LD); Great Atlas, R. Aït 

Messane, above Tinitive, c. 1500 m, 5 Jun 1926, H.Lindberg 3672 (LD); Kalaia, Mgoura, 21 Jun 1939, 

R.Maire & M.Weiller 388 (P); Oukaimeden to Tizin'ou Addi, P.H.Davis 55352 (BM, E); Haut Atlas, 

Ouarzazate, Msemrir, G.Bocquet 10885 (BM, ZT). 

Subsp. brachyanthus is more distinctive than the other subspecies and may merit recognition 

as a separate species. 

237

238 


187. Juncus oxycarpus E.Mey. ex Kunth, Enum. Pl. 3: 336 (1841) 

T: [Cape Provinces] Liesbeek R., C.H.Bergius; syn: B [destroyed]; Paarl, Berg Rivier, J.F.Drège a; syn: K, P. 

Juncus quartinianus A.Rich., Tent. Fl. Abyss. 2: 339 (1851). T: [Ethiopia] Shire [Chirϑ], R.Quartin-Dillon 

& A.Petit; holo: P; iso: K, L. 

Juncus brevistilus Buchenau, Monogr. Juncac. Cap 433 (1875) [Abh. Naturwiss. Vereine Bremen 4: 433 

(1875)]. T: Caput bonae spei, [collector not given] [label written by C.F.Hochstetter]; holo: n.v. [details 

depicted by F.G.P.Buchenau, Monogr. Junc. Cap, tab. 8, fig. 1–5]. 

Juncus gentilis N.E.Br., Kew Bull. 2: 83 (1914). T: Transvaal, Modderfontein, P.Conrath 1173; holo: K. 

Juncus oxycarpus var. microcephalus Adamson, J. Linn. Soc. Bot. 50: 13 (1935). T: numerous syntypes, 

e.g.: Cape, Riversdale, Muir 3385; syn: BOL; Grahamstown, R.A.Dyer 173; syn: BOL. 

Juncus suboxycarpus Adamson, J. Linn. Soc., Bot. 50: 14 (1935). T: Natal, Clairmont, R.Schlechter 3043; 

holo: BOL; iso: PRE. 

Juncus oxycarpus subsp. sparganioides Weim., Svensk Bot. Tidskr. 40: 166 (1946). T: Mt Kenya, Liki, 

11 Feb 1922, R.E.Fries & T.C.E.Fries 1477; holo: UPS; iso: LD. 

Juncus oxycarpus subsp. australis Weim., Svensk Bot. Tidskr. 40: 166 (1946), nom. inval.; probably meant as 

a name for the type subspecies. 

Illustrations: F.G.P.Buchenau, Monogr. Juncac. Cap, tab. 5 (1875); A.A.Obermeyer, in A.A.Obermeyer, 

J.Lewis & R.B.Faden, Fl. S. Afr. 4(2): 81, fig. 24 (1985). 

Maps: H.Weimarck, Svensk Bot. Tidskr. 40: 165, fig. 13 (1946); A.A.Obermeyer, in A.A.Obermeyer, J.Lewis 

& R.B.Faden, Fl. S. Afr. 4(2): 80, map 63 (1985) [both South Africa only]. 

Perennials, (25–) 30–70 (–80) cm tall, densely caespitose; rhizome short, horizontal, thick, shortnoded, 

in running water with nodal roots and innovations; stem terete. Cataphylls 0–2 (–3); 

basal leaves 1–2. Cauline leaves 2–3, terete, unitubular, distinctly perfectly septate, of 

variable length (8–50 cm), c. 1–2 mm wide; auricles obtuse, oblong-rounded, 0.8–2.0 mm 

long, yellowish-subcoriaceous to ±scarious. Lower bract leaf-like, 0.5–2.5 mm long, much 

shorter than inflorescence. Inflorescence compound to decompound, often divaricate, usually 

3–15 cm long, of 3–15 semiglobose to ±globose, c. 9–20-flowered heads of ±rigid branches. 

Capitulum bracts pale brownish to scarious, aristate. Tepals ±equal, lanceolate, c. 3.2–4.5 mm 

long, acuminate; central band greenish to stramineous-brown bordered red-brown to 

castaneous; margins broad, membranous. Stamens 3, less often to 6; anthers 0.5–0.8 mm 

long, shorter than 0.7–1.0 mm filaments; style 0.1–0.2 mm long (rarely to 0.4 mm); stigmas 

red, c. 1.5 mm long. Capsule oblong-ovoid, trigonous, unilocular, subobtuse, apiculate, rarely 

acute, c. 2.8–4.1 mm long, castaneous-brown, shining, subequalling perianth. Seeds ellipsoidovoid, 

c. 0.5–0.6 × 0.2–0.3 mm, apiculate, pale brown, longitudinally c. 15–20-striate, 

irregularly transversally reticulate; appendages absent. 

E & C part of Africa from Ethiopia, Eritrea and Somalia, through Tanzania, Zaire, Zambia, 

Angola, Zimbabwe and Botswana to South Africa. 23: BUR, RWA, ZAI. 24: ERI, ETH, 

SOM. 25: KEN, TAN. 26: ANG, MLW, ZAM, ZIM. 27: BOT, CPP, LES, NAT, TVL. 

Swamps, streambanks, seepage sites. Map 384. 

24. ERITREA: Saganeïti, Gorge de Degerra, 2200 m, G.A.Schweinfurth & D.Riva 892 (P). ETHIOPIA: Bale 

Province, Gurie, H.F.Mooney 7270 (K). SOMALIA: 'Somaliland', 10°57'N 48°38' E, J.G.B.Newbould 912 

(K) [longer auricles and rostrum suggest an intermediate position between J. fontanesii and 

J. oxycarpus]. 25. TANZANIA: Usambara, M.Batty 985 (P). 26. ANGOLA: Serra da Chella, Humpata, 

H.Humbert 16656 (P). ZAMBIA: Munshiwemba, Sep 1940, F.O.Stöhr 266 (BOL). ZIMBABWE: Harare 

[Salisbury], Nov 1919, F.Eyles 1891 (NBG-SAM). 27. CAPE PROVINCES: Elliott, Sow Veld, 

J.P.H.Acocks 20435 (K). LESOTHO: Underberg, Sehlabathebe National Park, Maal, F.K.Hoener 2117 (S). 

KWAZULU-NATAL: O.M.Hilliard & B.L.Burtt 17557 (E, K, NBG).. 

Plants from Eritrea and Ethiopia used to be referred to as J. fontanesii in the literature [e.g., 

J.G.Baker, in W.T.Thiselton-Dyer, Fl. Trop. Afr. 8: 94 (1902)], and indeed some specimens, 

otherwise matching J. oxycarpus have longer capsules and approach J. fontanesii subsp. 

pyramidatus. These plants and the ones of the latter taxon from SW Arabian peninsula need a 

new revision.


188. Juncus exsertus Buchenau, Monogr. Juncac. Cap 435 (1875) [Abh. Naturwiss. 

Vereine Bremen 4: 435 (1875)] 

T: [Cape Provinces, Swartkops River] Zwartkops Rivier, K.L.P.Zeyher 103; syn: B [destroyed after having 

been selected as type by R.S.Adamson, J. Linn. Soc. Bot. 50: 15 (1935)], BOL; Worcester, Waterfall, 

C.F.Ecklon & K.L.P.Zeyher [as Juncus punctorius 1. 11] p.p.; syn: B [destroyed], PRE; Zondagsrivier bei 

Graaff-Reinet [Sundays River at Graaff-Reinet], H.Bolus 188; syn: BOL; ‘Camdeboosberg, 4–5000 Fuss’, 

J.F.Drège [Juncus oxycarpus ‘c’]; syn: W [only!]. 

Juncus rostratus Buchenau, Monogr. Juncac. Cap 437 (1875) [Abh. Naturwiss. Vereine Bremen 4: 437 

(1875)]. T: [Cape Provinces, Swartkops River] Zwartkops Rivier, C.F.Ecklon & K.L.P.Zeyher; syn: S 

[possibly lecto, fide R.S.Adamson, J. Linn. Soc. Bot. 50: 17 (1935)]; Basche [Bashee R.], am Ufer des Flusses, 

J.F.Drége 4465; syn: K, G. 

Juncus exsertus subsp. lesuticus Burtt, Notes Roy. Bot. Gard. Edinburgh 45(2) (1988): 191 (1989). T: Cape, 

Barkly East, Witteberg, Ben McDhui, 8600 ft., 4 Feb 1983, O.M.Hilliard & B.L.Burtt 16435; holo: E; iso: NU, UPS. 

Illustration: F.Buchenau, Monogr. Juncac. Cap, tab. 5 (1875). 

Map: A.A.Obermeyer, in A.A.Obermeyer, J.Lewis & R.B.Faden, Fl. S. Afr. 4(2): 82, map 64 (1985). 

Perennials, usually 30–55 cm tall, caespitose; rhizome short, horizontal, with abbreviated 

internodes. Cataphylls 1–2, brownish; basal leaf usually single or absent; Cauline leaves 

2–3 (–4), terete, unitubular, distinctly septate, c. 5–16 cm long, 1.2–3.5 mm in diam.; 

auricles obtuse to rounded, 1–3 mm long, usually yellowish. Lower bract sheath-like, with 

blade short, 0.5–3 cm long. Inflorescence rich, decompound, much-branched, of (5–) 10 to 

many obconical to semiglobose, (2–) 3–7 (–15)-flowered heads 5–9 mm wide; capitulum 

bracts ovate, acute, scarious, brownish at base. Tepals ±equal, 2.4–3.5 mm long; outer tepals 

lanceolate-carinate, with a green and veined central band, dark castaneous-brown in upper 

part; inner tepals broadly lanceolate, acute to mucronate, with broad distinct membranous 

margins. Stamens (3–) 6; anthers 0.4–0.8 mm long; filaments 0.5–0.7 mm long; style c. 0.2 mm 

long; stigma reddish, 1.0–1.5 mm long. Capsule unilocular, narrowly trigonous-prismatic or 

oblong-trigonous, gradually narrowing in c. 0.3–0.5 mm rostrum, 2.8–3.8 mm long, 

castaneous-brown to dark brown, shining, much exceeding perianth. Seeds ellipsoid-ovoid, 

c. 0.5–0.6 × 0.2–0.3 mm, apiculate, pale brown, c. 15–20-striate longitudinally, transversely 

reticulate; appendages absent. 

Restricted to South Africa and Zimbabwe. 26: ZIM. 27: CPP, LES, NAM, NAT, OFS, SWZ, 

TVL. River banks, marshes. Map 385. 

27. CAPE PROVINCES: S Cape, Anysberg R., Vredeopstal, A.J.Martin 6 (NBG); Katberg, R.S.Adamson 305 

(BOL); Worcester, K.L.P.Zeyher s.n. (K); E Cape, Ibeka, 14 Jan 1895, R.Schlechter 6254 (WU). LESOTHO: 

Blue Mountain Pass, O.M.Hilliard & B.L.Burtt 17706 (E, K). KWAZULU-NATAL: Umtata, R.Schlechter 

6348 (WU). FREE STATE: Ladybrand, 1650 m, 27 Dec 1958, E.Werdermann & H.D.Oberdieck 1571 (K); 

Bloemfontein, A.Rehmann [Exs. Afr. Austr.] 3762 (K). NORTHERN PROVINCES: Transvaal, Heidelberg, 

towards Vereeniging R., A.Balsinhas 3366 (K); Johannesburg, R.Rand 1032 (BM). 

189. Juncus alpinoarticulatus Chaix, Pl. Vapinc. 74 (1785), & Chaix, in D.Villars, 

Hist. Pl. Dauph. 1: 378 (1786) 

Juncus alpinus Vill., Hist. Pl. Dauphiné. 2: 233 (1787), nom. illeg.; Juncus mucroniflorus Clairv., Man. 

Herbor. Suisse 104 (1811), nom. illeg.; Juncus acutiflorus var. alpinus [Vill.] Gaudin, Agrost. Helv. 2: 223 

(1811); Juncus fuscoater var. minor Spenn., Fl. Friburg. 1: 183 (1825); Juncus ustulatus β [unranked] alpinus 

(Gaudin) Gaudin, Fl. Helv. 2: 553 (1828); Juncus alpinus A. [unranked] mucroniflorus [Clairv.] Asch. & 

Graebn., Syn. Mitteleur. Fl. 2(2): 472 (1904), nom. inval.; Juncus alpinus subsp. australis Hyl., Uppsala Univ. 

Årsskr. 1945 (7): 107 (1945), nom. inval. T: Haller 1321 [V.A.Haller, Hist. Stirp. Helv. 2: no. 1321 

(1768)]; Ober Stafel, V.A.Haller 4; lecto: P-Haller, fide L.Hämet-Ahti, Ann. Bot. Fenn. 17: 342 (1980). 

Juncus sosnowskyi Novikov, Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 94(1): 110 (1989). 

T: [Georgia] Caucasus, Prov. Kutais, Svanethia, Comn. Mestia, in pascuis alpinis supra p. Mestia, 25 Jul 1910, 

D.Sosnowsky; holo: TBI. 

Maps: E.Hultén, Kongl. Svenska Vetenskapsakad. Handl., ser. 5, 8(5): map 88 (1962); E.Hultén, Fl. Alaska 

292 (1968); E.Hultén & M.Fries, Atlas N.Eur. Vasc. Pl. 1: 95, map 189 (1986). 

Perennials, 5– 60 (–80) cm tall, with a horizontally creeping c. 1–2 mm thick rhizome. Stems 

erect, with 1–2 basal sheaths or leaves and (1–) 2–3 (–5) cauline leaves. Leaves unitubular, 

perfectly septate, usually furrowed at base; auricles obtuse. Inflorescence variable in form and 

239

240 


size. Tepals (1.5–) 2–3 (–3.8) mm long, equal or subequal, obtuse; outer tepals often 

mucronate, with narrow scarious margins. Stamens 6, 1/2– 2/3 as long as tepals; anthers 0.3–0.7 mm 

long, 1/2– 2/3 times as long as (0.2–) 0.4–0.9 mm filaments; style 0.2–0.3 mm long; stigmas 

1.0–1.5 mm long. Capsule 2–3.3 (–3.8) mm long, equalling or exceeding perianth, obtuse, 

mucronate, shiny. Seeds 0.5–0.6 × 0.2–0.3 mm, ovoid or obliquely ovoid, reticulate; 


A species with a very wide distribution range in Europe, N Asia, Greenland and North 

America, and NW Africa. The species is very variable and the six subspecies recognized 

here cover most of the variation range, but further study is needed. In particular, plants from 

Spain, E Asia and China should be re-examined. The name Juncus sosnowskyi, according to 

its author, is close to J. alpinoarticulatus but may represent a hybrid between J. articulatus 

and the latter. 

1 Separate long-pedicellate flowers several to many in the inflorescence; 

their pedicels usually 2–6 mm long; if long-pedicellate flowers lacking 

then pedicels c. 2 mm long and flower heads few 

2 Capsule 3.0–3.5 mm long, exceeding to much exceeding perianth; 

tepals of ±equal length 

3 Pedicels of separate long-pedicellate flowers not exceeding 2 mm; 

flower heads usually 1–5 189c. subsp. alpestris 

3: Pedicels of separate long-pedicellate flowers 2–6 mm long; flower 

heads usually 10–25 189d. subsp. rariflorus 

2: Capsule to c. 2.5 mm long, equalling to slightly exceeding perianth; 

outer tepals conspicuously longer than the inner ones 189e. subsp. americanus 

1: Separate long-pedicellate flowers absent; pedicels usually to 1 mm 

4 Capsule usually acute, much exceeding perianth; tepals 1.5–2.0 mm long 189f. subsp. fuscescens 

4: Capsule usually obtuse, equalling to exceeding perianth; tepals usually 

2.0–2.5 mm long 

5 Pedicels up to 0.5 mm long; capsule ovoid; flower heads usually 20–30 189a. subsp. alpinoarticulatus 

5: Longest pedicels 0.5–1.0 mm; capsule oblong to ±cylindric; flower 

heads usually 5–10 189b. subsp. fischerianus 

189a. Juncus alpinoarticulatus Chaix subsp. alpinoarticulatus 

Juncus fuscoater Schreb.fuscoater Schreb., in A.F.Schweigger & F.Koerte, Fl. Erlang. 149 (1811); Juncus 

aquaticus subsp. fuscoater (Schreb.) Hagenb., Tent. Fl. Basil. 1: 327 (1821); Juncus alpinus var. fuscoater 

(Schreb.) Rchb., [sched.] Fl. Germ. Exs., no 159 (c. 1845), and Buchenau, Bot. Jahrb. Syst. 12: 374 (1890); 

Juncus alpinoarticulatus subsp. fuscoater (Schreb.) O.Schwarz, Mitt. Thüring. Bot. Ges., 1(1): 90 (1949). 

T: [Germany] Erlangen, J.C.D.Schreber; syn: W. 

Juncus ustulatus Hoppe, Anleit. Gräser 30 (1819). T: [Austria] prope Heiligenblut, D.H.Hoppe; syn: K, W. 

Juncus lampocarpus var. pygmaeus Salis-Marschl. [Flora 16 (2): 488 (1833), nom. inval.] ex Buchenau, Krit. 

Verz. Juncac. 65 (1880); Juncus alpinus II. [unranked] pygmaeus (Salis) Asch. & Graebn., Syn. Mitteleur. Fl. 

2 (2): 473 (1904). T: Corsica, Bastia, supra Corte, U.Salis-Marschlins; syn: n.v. 

Juncus fuscoater [Spielart] coarctatus G.Mey., Chloris Han. 566 (1836), nom. inval. 

Juncus lampocarpus var. microcephalus Peterm., Flora 27(1): 362 (1844). T: [Poland, Świnoujscie] 

Swinemünde, Winkler; syn: n.v. 

Juncus lampocarpus var. obtusiflorus Neilr., Fl. Wien 97 (1846). T: not designated. 

Juncus carpaticus Simonk., Termeszetrajzi. Füz. 10: 184 (1886); Juncus alpinus A. II. [unranked] carpaticus 

(Simonk.) Degen ex Asch. & Graebn., Syn. Mitteleur. Fl. 2 (2): 473 (1904), nom. illeg., non Racib. (1888); 

Juncus alpinus subsp. carpaticus (Simonk.) Borza, Fl. Romaniae Exs., n.v. [nom. illeg. if valid]; Juncus 

alpinus var. carpaticus (Simonk.) T.Georgiev & Kozhukhov., Fl. Bulg. 2: 167 (1965). T: numerous 

syntypes from Romania; citation: in alpis Bihariensibus (Cornul-muntyeluj, La-grope, Valye-csepilor), in 

alpibus Kiralyko et Bucsecs, nec non in alpibus Rodnensibus (Unoko). Transsilvania, in valle Csepilor alpium 

Bihariensium, 15 Jul 1882, L.Simkovics [=Simonkai]; syn: WU; [Romania] in turfosis secus rivulos alpium 

Bihariensium inter Paltinet et Pietra-tosa, 18 Jul 1879, L.Simonkai; syn: WU [both specimens originally named 

Juncus alpinus var. cuspidatus Simonkai, the latter later renamed Juncus carpaticus].


Juncus alpinus f. carpathicus Racib., Consp. Junc. Polon. 17 [=p. separ.] (1888). T: [Ukraine] Połonina 

Dzymbronia w Czarnej horze, A.Śleńdziński; holo: n.v. 

Juncus fuscoater var. microcarpus Brenner, Meddel. Soc. Fauna Fl. Fenn. 16: 55 (1889). T: [not indicated, 

authentic material not found in H]. 

Juncus fuscoater subsp. arthrophyllus Brenner, Meddel. Soc. Fauna Fl. Fenn. 16: 56 (1889); Juncus alpinus 

subsp. arthrophyllus (Brenner) Hyl., Nord. Kärlväxtfl. 1: 171 (1953), nom. inval.; Juncus alpinoarticulatus 

subsp. arthrophyllus (Brenner) Reichg., in C.J. van Oooststroom & al., Fl. Neerl. 1(6): 195 (1964). 

T: Finland, Nylandia, Kyrkslätt, Weikkola, 1886, M.M.W.Brenner; lecto: H, designated here by L.Hämet-Ahti. 

Juncus alpinus var. grandiflorus Beyer, Progr. Andr. Realgymn. Berlin 1891: 27 (1891), n.v. T: zwischen 

Revers und Ceres in den Grajischen Alpen, R.Beyer; syn: n.v. 

Juncus fuscoater f. subatricapillus Neuman, Sver. Fl. 657 (1901). T: not designated. 

Juncus carpaticus f. minor Zapał., Consp. Fl. Gallic. Crit. 1: 136 (1906). T: [Ukraine, the Carpathians] Pop 

Ivan, Kiputa, Wołoszczak; syn: n.v.; [Ukraine, the Carpathians] Dzembronia w krainie kosodrzewu Czarnej 

Hory [krummholz belt of Dzembronia, Czarna Gora Range], A.Ślendziński; syn: n.v. 

Juncus alpinus f. giganteus Erdner, Ber. Naturwiss. Vereins Schwaben u. Neuburg 39–40: 114–115 (1911). 

T: [Germany, Bavaria] Schwaben, Neuburg a. D.] Ufersand des Lechs [Lech R.] zwischen Feldheim und 

Marxheim, E.Erdner; syn: n.v. 

Juncus alpinus f. acicularis Lindq., Bot. Not. 1932: 354 (1932). T: not designated. 

Illustrations: T.Szynal & J.Mądalski, Atlas Fl. Pol. 2(2): 139a (1931); T.J.Reichgelt, Fl. Neerl. 1 (6): 193, fig. 

54 (1964); Ö.Nilsson & S.Snogerup, Bot. Not. 125: 134, fig. 71 (1972). 

Inflorescence narrow to broad in outline; main branches ±erect to patent; flower heads 

(6–) 25–30, (3–) 4–5 (–6.5) mm in diam., (2–) 3–5 (–10)-flowered; outermost flowers erect to 

patent; long-pedicellate separate flowers absent; longest pedicels c. 0.5 mm long. Tepals ±equal, 

all usually obtuse, c. (1.5–) 2.0–2.5 mm long. Capsule ovoid, usually obtuse, less often subacute, 

2.0–2.5 mm long, equalling or slightly exceeding perianth. 2n = 40, fide S.Snogerup, Bot. Not. 

111: 250 (1958); M.Zandee, Proc. Kon. Akad. Wet., ser. C, 84: 250 (1981). Fig. 106. 

Widely distributed in Europe, reaching the Caucasus and Turkey in the east. Probably 

introduced to West Siberia (Tomsk). Recorded from Morocco but material not seen. Spanish 

plants are similar to subsp. alpestris but not identical. 10: DEN, FIN, GRB, SWE. 11: AUT, 

CZE, GER, HUN, NET, POL, SWI. 12: FRA, SPA. 13: ALB, BUL, GRC, ITA, ROM, TUE, 

YUG. 14. BLR, BLT, RUC, RUN, RUS, RUW, UKR. 20: MOR. 30: wsb. 33: NCS, TCS. 34: 

TUR. Wet meadows, along streams, often calcicole but not always so. Map 386 (secondary 

occurrence in Siberia not shown). 

10. SWEDEN: Småland, Jönköping, Rocksjön, 23 Aug 1926, N.Hylander (UPS). 11. GERMANY: Berlin, 

Jungfernheide, H.G.L.Reichenbach [Fl. Germ. Exs.] 159 (K, PRC). SWITZERLAND: Chabrey, 430 m, 

Thiébaut [Exs. Genav.] 64 (LD). 12. FRANCE: Strasbourg, Plobsheim, 20 Jul 1971, Kapp [Soc. Éch. Pl. 

Vasc. Bas. Médit.] 6343 (LD, P). SPAIN: Asturias, Somiedo, lago Cerveriz, 28 Jul 1976, J.A.Fernandez 

Prieto (FCO9839); Granada, Sierra Nevada, Prados de Otero, 7 Jul 1981, M.C.Fernández-Carvajal (FCO 

19923); Huesca, Valle de Ordesa, 19 Jul 1976, F.Navarro, T.E.Díaz & M.C.Fernández-Carvajal (FCO6438); 

León, Macizo Central de los Picos de Europa, Liordes, 1890 m, 25 Aug 1983, H.S.Nava (FCO13895); Lérida, 

Boi, Estany Negre, 6 Jul 1958, P.Montserrat (JACA). 14. NORTHWEST EUROPEAN RUSSIA: Sankt 

Peterburg, Pavlovsk, D.Litvinov [Gerb. Fl. SSSR] 6972 (W). 

Spanish populations of Juncus alpinoarticulatus are tentatively included in the type 

subspecies. However, they are distinct from the most common morphotypes in several 

respects, and may prove to represent a separate taxon. 

189b. Juncus alpinoarticulatus subsp. fischerianus (Turcz. ex V.I.Krecz.) Hämet-Ahti, 

Mem. Soc. Fauna Fl. Fenn. 56: 97 (1980) 

Juncus fischerianus Turcz. ex V.I.Krecz., Fl. Transbaic. 2: 142 (1931). T: [Russia] Siberia, ad Baicalem 

prope monasterium Possolsky, 1831, P.K.N.S.Turczaninov; lecto: LE, fide L.Hämet-Ahti, Mem. Soc. Fauna Fl. 

Fenn. 56: 97 (1980); iso: G. 

Juncus alpinus var. macrocephalus Hyl., Bot. Not. 1953: 353 (1953). T: Finland, Savonia borealis, 

Maaninka, 1910, O.Kyyhkynen; lecto: H, fide L.Hämet-Ahti, Mem. Soc. Fauna Fl. Fenn. 56: 97 (1980). 

Illustration: L.Hämet-Ahti, Ann. Bot. Fenn. 23: 277 (1986). 

241

242 


Figure 106. Juncus alpinoarticulatus subsp. alpinoarticulatus. A, habit; B, inflorescence; 

C, tepals and stamens; D, perianth and capsule. Scale bars: A, B = 7 cm; C = 1 mm; D = 1.3 mm. 

Drawn by Ö.Nilsson. Reproduced with permission, from Ö.Nilsson & S.Snogerup, Bot. Not. 

125: 134, fig. 71 (1972). © Lund Botanical Society.


Inflorescence broad in outline; main branches usually broadly patent; flower heads usually 

5–10, 7–8 (–10) mm in diam., (5–) 8–10 (–12)-flowered; outermost flowers patent; longpedicellate 

separate flowers absent; pedicels to 1 mm long. Tepals ±equal, all usually obtuse, 

c. 2.0–2.5 mm long. Capsule oblong to cylindric, obtuse, c. (2.5–) 3.0 mm long, exceeding 

perianth. 

The subspecies occupies a large range from E Scandinavia, through N European Russia, 

Siberia, to the Far East; it is also known from China (Hebei). Reported from Sakhalin but 

material not seen. Reports of the occurrence of J. alpinus in Middle Asia (Kazakhstan) may 

refer to subsp. fischerianus but material was not seen during this study. 10: FIN. 14: BLT, 

RUN, RUW. 30: ALT, BRY, CTA, IRK, KRA, WSB, YAK. 31: KHA. 32: KAZ [?]. 36: 

CHN. 37: MON. Lake shores, along streams, wet ditches, less often roadsides. Map 387. 

10. FINLAND: Savonia borealis, Joroinen, Kotkalahti, 1950, O.Tiitinen (H). 14. ESTONIA: Peipsjärv, 

Mustvee, 1964, Saare (TAA). NORTH EUROPEAN RUSSIA: Vologda, Tetshara R., 1907, R.R.Pohle (LE); 

Komi Republic, Syktyvkar, 1967, H.Aasamaa 11261 (H). 30. ALTAI: Kurai R., G.P.Sumnevich s.n. (LE). 

IRKUTSK: Irkutsk, Bratsk–Ust' Kut, 1922, Popov (LE). KRASNOYARSK: Evenki Region, Tunguzka– 

Chunya Rayon, 1968, Sokolova (LE). WEST SIBERIA: Omsk, Tara Distr., 1912, Birevich 789 (LE). 

36. CHINA NORTH-CENTRAL: Hebei, Ying Ho Chuang Ho R. by Yang Shu Pei, 1891, Putyata (LE). 

189c. Juncus alpinoarticulatus subsp. alpestris (Hartm.) Hämet-Ahti, Ann. Bot. Fenn. 

17: 342 (1980) 

Juncus alpinus subsp. alpestris (Hartm.) Á.Löve & D.Löve, Rep. Dept. Agric. Univ. Inst. Appl. Sci. 

(Reykjavik) ser. B, 3: 23 (1948); Juncus alpestris Hartm., Handb. Skand. Fl. 141 (1820); Juncus ustulatus var. 

alpestris (Hartm.) Hartm., Handb. Skand. Fl., ed. 2, 94 (1832); Juncus alpinus var. alpestris (Hartm.) 

C.Hartm., Handb. Skand. Fl., ed. 11: 422 (1879); Juncus nodulosus var. alpestris (Hartm.) Shljakov, in 

O.I.Kuzeneva & al., Fl. Murmansk. Oblasti 2: 172 (1954), nom. inval. T: [Sweden, Lule Lappmark] 

Quickjock, L.L.Laestadius; syn: UPS. 

Juncus rariflorus var. obtusatus Hartm., Handb. Skand. Fl. 141 (1820); Juncus alpinus f. obtusatus (Hartm.) 

Lindq., Bot. Not. 1932: 354 (1932). T: [Sweden, Lule Lappmark] Quickjock, L.L.Laestadius; lecto: UPS, 

designated here. 

Juncus nodulosus var. biceps Laest. ex Wahlenb., Fl. Suec., ed 2: 1090 (1833). T: [Sweden/Finland] 

Karesuvando, L.L.Laestadius; syn: LD, UPS. 

Juncus nodulosus var. uniceps Laest. ex Wahlenb., Fl. Suec. ed 2, 1090 (1833); Juncus alpinus γ [unranked] 

uniceps (Wahlenb.) Hartm., Handb. Skand. Fl. ed. 7, 240 (1858); Juncus nodulosus f. uniceps (Laest.) 

Shljakov, in O.I.Kuzeneva & al., Fl. Murmansk. Oblasti 2: 172 (1954), nom. inval. T: [Sweden/Finland] 

Karesuvando [Karesuando/Kaaresuvanto], L.L.Laestadius; syn: LD, UPS. 

Illustration: Ö.Nilsson & S.Snogerup, Bot. Not. 125: 136, fig. 73 (1972). 

Maps: B.Lindquist, Bot. Not. 1932: 345 (1932); B.Lindquist, Acta Phytogeogr. Suec. 13: 125 (1940). 

Plants 5–20 cm tall. Rhizome creeping, with internodes 1–10 mm long. Inflorescence narrow 

in outline; main branches erect; heads 1–5 (–7), 2–5 (–7) mm wide, each with (3–) 4–9 dark 

brown to castaneous-coloured flowers; outermost flowers of heads erect, long-pedicellate; 

separate flowers few to absent with pedicels to 2 mm long. Tepals ±equal 2.0–3.0 mm long, 

all broadly obtuse; outer tepals often mucronate. Capsule ellipsoid-ovoid, (2.5–) 3.0–3.5 mm 

long, ±exceeding perianth, obtuse to ±subacute, shortly mucronate, very dark castaneous to 

blackish, shiny. 

Iceland, mountains and Arctic part of Scandinavia, and N Russia. 10: FIN, ICE, NOR, SWE. 

14: RUN. Streamsides, acid fens dominated by sedges, on peaty soils. Map 388. 

10. FINLAND: Kuttanen, Kaaresuvando, 7 Aug 1907, J.Montell (UPS). 

189d. Juncus alpinoarticulatus subsp. rariflorus (Hartm.) Holub, Folia Geobot. 

Phytotax. 23: 413 (1988) 

Juncus rariflorus Hartm., Handb. Skand. Fl. 141 (1820); Juncus ustulatus β [unranked] rariflorus (Hartm.) 

Lange, Haandb. Danske Fl. 213 (1851); Juncus alpinus var. rariflorus (Hartm.) Hartm., Handb. Skand. Fl. ed. 

7, 240 (1858); Juncus alpinus subsp. rariflorus (Hartm.) K.Richt., Pl. Eur. 1: 181 (1890). T: [Sweden, 

Stockholm] Holmiae ad ripas lacuum, C.J.Hartman; lecto: UPS, designated here. 

243

244 


Juncus nodulosus Wahlenb., Fl. Upsal. 114 (1820); Juncus alpinus subsp. nodulosus (Wahlenb.) Lindm., 

Svensk Fanerogamfl. 155 (1918); Juncus alpinoarticulatus subsp. nodulosus (Wahlenb.) Hämet-Ahti, Ann. 

Bot. Fenn. 17: 342 (1980). T: [Sweden] Uppsala, på Mälarns strand bortom Vårdsätra, 25 Aug 1819, 

G.Wahlenberg; lecto: UPS, fide R.Moberg & Ö.Nilsson, Nordic J. Bot. 11: 297 (1991) [further Wahlenberg 

material in S, UPS]. 

Juncus nodulosus f. dissolutus Lindm., Svensk. Fanerogamfl. 165 (1918). T: not designated. 

Juncus marshallii Pugsley, J. Bot. 69: 279, 282 (1931); Juncus alpinus var. marshallii (Pugsley) Lindq., Bot. 

Not. 1932: 355 (1932); Juncus nodulosus var. marshallii (Pugsley) P.W.Richards, in A.R.Clapham, T.G.Tutin 

& E.F.Warburg, Fl. Brit. Isles 1251 (1952); Juncus alpinoarticulatus var. marshallii (Pugsley) P.D.Sell, in 

P.D.Sell & G.Murrell, Fl. Gr. Brit. Irel. 5: 363 (1996). T: Scotland, Loch Ussie near Dingwall, E Ross, 

E.S.Marshall; holo: CGE; iso: E, photo PRA. 

Juncus alpinus f. grandiceps Lindq., Bot. Not. 1932: 354 (1932). T: not designated. 

Juncus alpinus f. pygmaeus Lindq., Bot. Not. 1932: 355 (1932), nom. illeg., non Buchenau. T: not designated. 


Map: B.Lindquist, Acta Phytogeogr. Suec. 13: 124, 125 (1940). 

Inflorescence narrow in outline; main branches ±erect; heads (5–) 10–25, 2–5 mm in diam., 

(1–) 3–5 (–7)-flowered; outermost flowers in heads erect; long-pedicellate separate flowers 

present, several to many, with pedicels 4–6 mm long. Tepals ±equal, 2.5–3 mm long; outer 

tepals usually subacute; inner tepals obtuse. Capsule subcylindric to narrowly ovoid, 

subobtuse to acute, 3.0–3.5 mm long, much exceeding perianth. 2n = 40, fide S.Snogerup, 

Bot. Not. 111: 250 (1958). Fig. 107. 

Widely distributed from Iceland and Scandinavia eastwards to N European Russia and the 

westernmost northern Siberia. It is probably found at isolated localities in SW Siberia and 

Yakutia but the records require confirmation (material not seen). Reports from the Far East 

refer to subsp. americanus. 10: DEN, FIN, GRB, ICE, NOR, SWE. 14: RUN. 30: KRA?, 

TVA?, WSB, YAK?. Wet meadows, sandy or gravelly shores of ponds and streams. 

Map 389 (uncertain Asian localities not shown). 

10. FINLAND: Hämeenlinna, Luolaja, R.Lampinen 8972 (W); Oulun Pohjanmaa, Haukipudas, Loukkojärvi, 

Martimojoki, 12 Jul 1984, M.Ohenoja (LD, OULU). NORWAY: Vest-Agder, Lista, S of Farsund, 

Lomsesanda, S.Snogerup & B.Snogerup 9542 (LD); Opland, Snertingdal, near the farm of Granum, 20 Jul 

1949, K.H.Mattisson (LD). SWEDEN: Skåne, Ringsjön, 15 Aug 1885, Nilsson (S); Skåne, Stehag, Lake 

Ringsjön, 1 Aug 1920, O.R.Holmberg (LD); Skåne, Osby parish, at Lake Osbysjön, 3 Aug 1959, S.Snogerup 

(LD); Dalarna, Grangärde, Norrbo, 22 Jul 1972, A.Lundegren (LD); Ångermanland, Högsjö, Vedastrand, 28 

Aug 1981, E.Evers (LD); Norrbotten, Överluleå parish, S Brotjärn, c. 5 km SW of Boden Station, S.Snogerup 

775 (LD, 2n = 40). 

189e. Juncus alpinoarticulatus subsp. americanus (Farw.) Hämet-Ahti, Ann. Bot. 

Fenn. 23: 277 (1986) 

Juncus alpinus var. americanus Farw., Amer. Midl. Naturalist 11: 74 (1928). T: USA, Michigan, Keweenaw 

Co, Clifton, 1894, O.A.Farwell 837a; holo: BLH. 

Juncus affinis R.Br., Addenda, in J.Richardson, Bot. App. Narr. Journ. Franklin 735 (1823), nom. illeg., non 

Gaudin (1811); Juncus richardsonianus Schult. & Schult.f., Syst. Veg. 7(1): 201 (1829). T: [Canada] 

wooded country from latitude 54° to 64° north; syn: BM; Bear Lake, Dr. [J.]Richardson; syn: K. 

Juncus alpinus var. insignis Fr. ex Buchenau, Bot. Jahrb. Syst. 12: 374 (1890). T: numerous syntypes from 

North America, incl. Rocky Mtns, Canby 336; syn: n.v.; Detroit, J.M.Bigelow [G.Engelmann, Herb. Junc. 

Bor.-Amer. Norm.] 51; syn: MO, PR, S. 

Illustrations: L.Hämet-Ahti, Ann. Bot. Fenn. 23: 277 (1986); S.E.Clemants, Junc. New York State 42 (1990). 

Inflorescence narrow in outline; main branches erect; heads (3–) 10–20 (–40), usually 3–5 mm 

in diam., usually 3–5 (–12)-flowered; outermost flowers erect; separate long-pedicellate 

flowers present, few to many, with pedicels 2–4 mm long. Tepals unequal; outer tepals 

longer, c. 2.2–2.8 mm long, subacute; inner tepals obtuse. Capsule cylindric-oblong, obtuse, 

c. 2.5 mm long, equalling to slightly exceeding perianth. 2n = 40, fide N.A.Harriman & 

D.Redmond, Rhodora 78: 733 (1976), as J. alpinus. 

Widespread in the northern United States and in Canada and in the Russian Far East 

(Kamchatka, Chukotka). 31: KAM, KHA, MAG. 70: ASK, GNL, NWT, YUK. 71: ABT, BRC,


Figure 107. Juncus alpinoarticulatus subsp. rariflorus. A, habit; B, C, details of 

inflorescence; D, tepals and stamens; E, perianth and capsule. Scale bars: A = 5 cm; B, C = 1 cm; 

D = 1 mm; E = 1 mm. Drawn by Ö.Nilsson. Reproduced with permission, from Ö.Nilsson 

& S.Snogerup, Bot. Not. 125: 135, fig. 72 (1972). © Lund Botanical Society. 

245

246 


MAN, SAS. 72: LAB, NBR, NFL, NSC, ONT, PEI, QUE. 73: COL, IDA, MNT, WAS. 74: ILL, 

IOW, MIN, MSO, NDA, NEB, SDA, WIS. 75: INI, MAI, MAS, MIC, OHI, PEN, VER. 76: 

UTA. Wet meadows, sandy or gravelly shores, fens, clayey pools, often calcicole. Map 390. 

70. ALASKA: Shumagin Is., Popoff Is., E.Hultén 7719 (S); Skagway, J.P.Anderson 1839 (S). 

GREENLAND: Majut, 3 Sep 1962, C.Hansen & al. [Pl. Vasc. Groenl. Exs.] 138 (UPS). 71. BRITISH 

COLUMBIA: Saltspring Is., Saint Marie Lake, A.Ceska & O.Ceska s.n. (PRA, V). 72. ONTARIO: Thunder 

Bay District, Sibley Provincial Park, J.A.Bailey 1747 (PRA); Ontario, Thunder Bay District, Selim Hill, 

J.A.Bailey 1720 (PRA). QUEBEC: Carleton, J.F.Collins, M.L.Fernald & A.S.Pease [Pl. Exs. Gray.] 187 

(S); Bonaventure, Carleton, L.Cinq-Mars L-74 (LD). 75. MICHIGAN: Detroit, J.M.Bigelow [G.Engelmann, 

Herb. Junc. Bor.-Amer. Norm.] 51 (PR, S). 

Plants from S Greenland are ±intermediate between subsp. nodulosus and subsp. americanus 

but closer to the latter [see C.Hansen & al. 138 (UPS)]. Similarly intermediate plants are 

known from Inner Mongolia [E of Dojen, J.Eriksson 39 (S)] and some Siberian localities. 

The distribution of subsp. americanus requires further study. 

189f. Juncus alpinoarticulatus subsp. fuscescens (Fernald) Hämet-Ahti, Ann. Bot. 

Fenn. 23: 280 (1986) 

Juncus alpinus var. fuscescens Fernald, Rhodora 10: 48 (1908). T: New York, Cayuga Marshes, 1885, 

W.R.Dudley 137; lecto: GH, fide L.Hämet-Ahti, Ann. Bot. Fenn. 23: 280 (1986). 

Inflorescence broad in outline; main branches erecto-patent to patent; heads 10–20, usually 

4–6 mm in diam., 6–10-flowered; outermost flowers in heads patent; long-pedicellate separate 

flowers absent; pedicels to 0.5 mm long. Tepals ±equal, 1.5–2.0 mm long, all obtuse. 

Capsule oblong-ovoid, c. 2.5 mm long, acute, much exceeding perianth. 

Restricted to the southern vicinity of the Great Lakes, from New York to Minnesota. 74: MIN, 

WIS. 75: MIC, NWY. Wet meadows, sandy shores. Map 391. 

74. MINNESOTA: Ramsay Co., 1922, Rosendahl & al. 5184 (GH). WISCONSIN: Adams Co., 1932, 

N.C.Fassett & al. 14463 (GH); Marquette Co., 1968, N.A.Harriman 4032 (OSH). 75. MICHIGAN: Grand 

Traverse Co., 1934, F.J.Hermann 6288 (GH). NEW YORK: Cayuga Co., Lake Cayuga, K.M.Wiegand & 

E.J.Palmer 3772 (GH); Oswego Co., N of Selkirk, 1906, W.W.Rowlee s.n. (GH); Oswego Co., N of Selkirk, 

1906, W.W.Rowlee s.n. (GH). 

190. Juncus requienii Parl., Fl. Ital. 2: 346 (1852) 

Juncus alpinus var. requienii (Parl.) Husn., Joncées 15 (1908); Juncus alpinus subsp. requienii (Parl.) Nyman, 

Consp. Fl. Eur. 748 (1882). T: [Corse] in Corsica nel Monte Coscione, E.Requien; holo: FI, n.v. 

Perennials, 3–7 (–10) cm tall; rhizome creeping, branched, long-noded; stolons short. 

Cataphylls 1–2 (–3). Cauline leaves 2–3, unitubular, perfectly septate, to 5 cm long, 0.5–0.8 mm 

in diam.; apex ±acute; sheaths with membranous margins; auricles scarious to membranous, 

to 0.5 mm long. Lower bract ±leaf-like, to c. 2.5 cm long, or pale brown to castaneous, 

c. 0.5–1 cm long, shorter than inflorescence or longer when inflorescence reduced to a single 

head. Inflorescence a single terminal head, or of 2–6 obconical to semiglobose (2–) 3–4 (–7)flowered 

dark heads. Tepals ±ovate, subequal, 1.7–2.3 mm long; outer tepals oblong-ovate, 

mucronate, dark castaneous-brown or with a greenish central band; inner tepals oblong, 

obtuse, often involute, with margins broad and membranous. Stamens 6; anthers c. 0.6–0.9 mm 

long; filaments 0.5–0.8 mm long; style c. 0.5 mm long; stigmas to 1 mm long. Capsule 

unilocular, oblong-ovoid, trigonous, ±obtuse, subabruptly or gradually contracted into a 

0.2–0.3 mm rostrum, c. 2.2–2.6 mm long, pale brown to brown, shining, distinctly longer than 

perianth. Seeds ±ellipsoid, apiculate, c. 0.45–0.50 × 0.2–0.25 mm, pale brown, longitudinally 

c. 18–24-striate, finely reticulate; appendages absent. 

Endemic to Corsica. 12: COR. Fens and along streams, 1000–2300 m. Map 392. 

12. CORSE: Monte Renoso, 24 Jul 1901, R.Rotgés 458 (P); Palneca, 1200 m, 24 Jul 1901, R.Rotgés s.n. (P); 

Monte Renoso, Lac de Vitalacca, P.Mabille [Herb. Cors.] 400 (P, WU); Monte Coscione, 9 Aug 1879, 

E.Reverchon (LD).


191. Juncus anceps Laharpe, Essai Monogr. Jonc. 38 (1825) 

Juncus anceps var. genuinus Buchenau, Ber. Deutsch. Bot. Ges. 1: 493 (1883), nom. inval.; Juncus 

atricapillus subsp. anceps (Laharpe) K.Richt., Pl. Eur. 1: 181 (1890), nom. inval. T: Montpellier, 

P.Salzmann; syn: K; Mans, N.H.F.Desportes; syn: K; Bayonne, [collector not given]; syn: n.v. 

Juncus atricapillus Drejer [Bot. Tidsskr. 2: 182 (1838), here a plant from Jylland with dark flowers is 

mentioned under the name, which might have constituted a valid publication] ex Lange, Haandb. Danske Fl. 

212 (1851); Juncus anceps var. atricapillus (Drejer ex Lange) Buchenau, Ber. Deutsch. Bot. Ges. 1: 493 

(1883); Juncus anceps Rasse B. atricapillus (Drejer) Asch. & Graebn., Syn. Mitteleur. Fl. 2 (2): 475 (1904); 

Juncus alpinoarticulatus subsp. atricapillus (Drejer ex Lange) Reichg., in C.J. van Oooststroom & al., Fl. 

Neerl. 1(6): 193 (1964). T: [Denmark; citation: fra Skagen (Drej.) til Eidersted] Skagen, S.T.N.Drejer; syn: 

C, n.v., UPS. 

Juncus anceps var. sparsiflorus Lange, Haandb. Danske Fl., ed. 2, 237 (1856). T: not designated. 

Juncus anceps var. congestus Lange, Haandb. Danske Fl., ed. 2, 237 (1856). T: not designated. 

Juncus anceps subsp. herzegovinus Sagorski, Mitth. Thüring. Bot. Vereins 16: 36 (1901); Juncus anceps 

f. herzegovinus (Sagorski) Lindq., Bot. Not. 1932: 351 (1932). T: Hercegovina, Žiljevo, Nevesinsko Polje, 

29 Jul 1889, S.Murbeck; syn: UPS; Nevesinsko Polje, 2 Aug 1889, S.Murbeck; syn: LD. 

Juncus anceps f. pumilus Lindq., Bot. Not. 1932: 352 (1932). T: not designated. 

Juncus anceps f. coarctatus Lindq., Bot. Not. 1932: 351 (1932). T: not designated. 

Illustrations: T.J.Reichgelt, in C.J. van Ooststroom & al., Fl. Neerl. 1 (6): 193, fig. 54 (1964); Ö.Nilsson & 

S.Snogerup, Bot. Not. 125: 133, fig. 70 (1972). 

Map: E.Hultén & M.Fries, Atlas N.Eur. Vasc. Pl. 1: 95, map 190 (1986). 

Perennials, 20–60 cm tall; rhizome creeping, sparsely branched, with internodes of variable 

length. Cataphylls 0–2. Cauline leaves 3–5, unitubular, perfectly septate, 5–25 cm long, 

c. 1–2 mm in diam.; sheaths with distinct margins usually c. 1 mm wide; auricles 

subcoriaceous, obtuse scarious to brownish, c. 0.5–1.5 mm long. Lower bract usually short, 

sheath-like, with an awn-like blade, rarely leaf-like, much shorter than inflorescence. 

Inflorescence decompound, often divided in two discrete parts, of (25–) 50–80 semiglobose 

to obconical, (2–) 3–4 (–8)-flowered ±evenly spaced heads; primary branches ±erect; 

secondary branches erecto-patent. Tepals ±equal, ovate to oblong-ovate, 2.0–2.7 mm long; 

outer tepals ±acute, shortly mucronate; inner tepals broadly obtuse, usually dark brown to 

deep castaneous, with margins broad and scarious. Stamens 6; anthers 0.7–1 mm long; 

filaments 0.7–0.9 mm long; style 0.5–0.8 mm long; stigmas 1.0–1.5 mm long. Capsule 

unilocular, broadly ovoid-trigonous, usually subabruptly narrowing into a 0.3–0.5 mm 

mucro, 2.4–3.2 mm long, dark brown to deep castaneous, slightly exceeding perianth. Seeds 

ovoid-ellipsoid, c. 0.5–0.6 × 0.2–0.25 mm, apiculate, pale brown, longitudinally c. 25-striate, 

finely transversely reticulate; appendages absent. 2n = 40, fide S.Snogerup, in Ö.Nilsson 

& S.Snogerup, Bot. Not. 125: 132 (1972); M.Zandee, Proc. Kon. Ned. Akad. Wetensch., C, 

84: 250 (1981). Fig. 108. 

Mostly coastal areas in W, SW and S Europe, NW Africa. Reported to occur in Spain but no 

material was seen from the Iberian Peninsula, and the occurrence is doubtful. 10: DEN, fin, 

SWE. 11: BGM, GER, NET. 12: COR, FRA, SAR. 13: ALB, GRC, ITA, YUC. 20: ALG, 

MOR, TUN. Coastal sandy habitats, open sites in heathlands and fens. Map 393. 

10. DENMARK: Jutland, Skagen, Aug 1897, M.Lorentzen [Dörfler, Herb. Norm.]3856 (W, PR); Thyborøn, 

P.Holm & al. [Fl. Jutl. Exs.] 499 (C, W). 11. GERMANY: Juist Is., 5 Aug 1900, F.Buchenau & O.Lange 

[A.Kneucker, Cyper. Junc. Exs.] 53 (PR, PRA). THE NETHERLANDS: Terschelling, Boschplaat, 

K.U.Kramer & Lindeman [Fl. Neerl. Exs.] 222 (K). 12. FRANCE: Hérault, Palavas, 24 Jul 1899, G.Mandon 

(LD); Charente-Inférieure, Marais d'Anais, J.Foucaud [C.Magnier, Pl. Sel. Exs.] 136 (P). 13. ITALY: 

Toscana, Viareggio, 8 Aug 1869, E.Levier (LD); Toscana, Cascine di Firenze, Arno R., A.Contardo 

[Pl.Ital.Exs.] 25 (W). YUGOSLAVIA: Herzegovina, Kifinoselo, Jul 1902, E.Sagorski (LD); Nevesinsko 

Polje, 850 m, 10 Jul, E.Sagorski (W); Montenegro, Volujačko jezero, 8 Aug 1925, K.Maly (W). 

20. TUNISIA: Bou-Merchen, 27 Jun 1931, R.Maire (P). 

192. Juncus articulatus L. Sp. Pl. 327 (1753) 

Juncus articulatus [var.] aquaticus L., Syst. Nat., ed. 12, 250 (1767); Juncus aquaticus (L.) All., Fl. Pedem. 2: 

217 (1785); Juncus lampocarpus f. aquaticus (L.) Neuman, Sver. Fl. 656 (1901), nom. inval. 

247

248 


Figure 108. Juncus anceps. A, habit; B, condensed inflorescence; C, tepals and stamens; 

D, E, perianth and capsule. Scale bars: A, B = 3 cm; C = 1 mm; D, E = 1 mm. Drawn by 

Ö.Nilsson. Reproduced with permission, from Ö.Nilsson & S.Snogerup, Bot. Not. 125: 133, 

fig. 70 (1972). © Lund Botanical Society.


T: In Europae aquosis; C.Linnaeus; syn: LINN449/19 [recommended as lectotype by S.Snogerup, in A.Strid & 

K.Tan, Mountain Fl. Greece 2: 738 (1991), but further study needed]. 

Juncus tatewakii Satake, J. Jap. Bot. 14: 257 (1938); Juncus articulatus subsp. tatewakii (Satake) Vorosch., 

[Opred. Rast. Sov. Dal. Vost. 148 (1982), nom. inval.] in A.K.Skvortsov, Florist. Issled. Raz. Raion. SSSR 157 

(1985). T: Kuril Is., Insula Kunasiri, Itibisinai, 20 Aug 1936, M.Tatewaki 25464; holo: SAP; iso: TI. 

Perennials, (5–) 13–40 (–70) cm tall, caespitose or with a short-creeping rhizome, or at times 

with stems submerged or buried in sand and rooting at proximal nodes. Stems erect or 

ascending or rarely floating, with 0–2 greenish to brownish basal sheaths and 3–7 cauline 

leaves. Leaves unitubular, perfectly septate, terete or somewhat compressed; auricles scarious, 

obtuse, to c. 1 mm long. Lower bract leaf-like, usually to 4 cm long, shorter than 

inflorescence. Stamens 6, ( 1/3–) 1/2– 3/4 as long as tepals. Capsule unilocular, shortly mucronate, 

varying in colour. Seeds ovoid, 0.45–0.6 × c. 0.25 mm, pale brown, c. 25-striate, reticulate. 

Europe; Asia; N. Africa; North America; also in S. Africa, Australia and New Zealand 

(probably introduced). Various moist to wet places in natural and man-made vegetation, 

margins of watercourses, usually on acid soils. 

An extremely variable species. The variation is difficult to describe in terms of infraspecific 

taxa but two subspecies may be recognized; other similarly distinct variants may deserve a 

subspecific or varietal status but further study is needed. 

Plants pale green; tepals green, usually 2.4–2.8 mm long (inner tepals 

sometimes slightly longer), ±equal; capsule with rostrum 0.3–0.4 (–0.5) mm 

long; anthers conspicuously shorter than filaments 192b. subsp. limosus 

Plants mid-green; tepals with green central band bordered brown, or wholly 

brown, usually 2.2–3.5 mm long (outer tepals sometimes slightly longer), 

±equal; capsule with rostrum to c. 0.3 mm long; anthers equalling to slightly 

longer than filaments 192a. subsp. articulatus 

192a. Juncus articulatus L. subsp. articulatus 

Juncus geniculatus Schrank, Baier. Fl. 1: 613 (1789). T: [Germany, Bavaria; citation: Am Geradensteine im 

Gebirge bei Steingaden]; [locality not given], F.Schrank; syn: W. 

Juncus stolonifer Wohll., Suppl. Leyss. Fl. Halens. 13 (1796), n.v.; Juncus lampocarpus [unranked] stolonifer 

(Wohll.) Asch. & Graebn., Syn. Mitteleur. Fl. 2(2): 479 (1904); Juncus articulatus var. stolonifer (Wohll.) 

House, New York State Mus. Bull. 254: 213 (1924); Juncus articulatus f. stolonifer (Wohll.) Raymond, 

Naturaliste Canad. 77: 70 (1950). T: n.v. 

Juncus lampocarpus Ehrh. ex Hoffm., Deutschl. Fl., ed. 2, 1: 166 (1800); Juncus adscendens Host, Icon. 

Descr. Gram. Austriac. 3: 58 (1805), nom. illeg.; Juncus lampocarpus var. adscendens [Host] Čelak., Prodr. 

Fl. Böhmen. 1: 81 (1867); Phylloschoenus lampocarpus (Ehrh. ex Hoffm.) Fourr., Ann. Soc. Linn. Lyon, n. 

ser., 17: 171 (1869), nom. inval.; Juncus articulatus var. adscendens [Host] Beck, Fl. Nieder-Österreich 1: 

155 (1890); Juncus lampocarpus a. [unranked] eulampocarpus Asch. & Graebn., Syn. Mitteleur. Fl. 2 (2): 477 

(1904), nom. inval. T: [citation: Ehrhart gram. 126; F.Ehrhart, Calam. Tripet. Exs., no. 126 (1794–1795)]; 

[Sweden, Uppsala] Upsaliae, F.Ehrhart [Calam. Tripet. Exs.] 126; lecto: LINN-SMITH43, designated here 

[epithet often spelled as lamprocarpus]. 

Juncus aristiflorus Clairv., Man. Herbor. Suisse 104 (1811). T: Haller 1322 [V.A.Haller, Hist. Stirp. Helv. 

2: no. 1322 (1768)]; lecto: P-Haller, designated here. 

Juncus affinis Gaudin, Agrost. Helv. 2: 224 (1811); Juncus lampocarpus δ [unranked] affinis (Gaudin) 

Gaudin, Fl. Helv. 2: 552 (1828). T: [Switzerland] icon in C.Bauhin, Prodr. 12, fig. 2; syn.; 

[Switzerland/France; citation: aux Grangettes ad caput Lemani, 1801 & 1810, J.F.Gaudin] aux près de 

Villeneuve, Vaud, Gaudin; syn: K [material sent to J.Gay in 1803]; Grangettes, J.F.Gaudin; syn: K [material 

sent to J.Gay in 1810]. [Additional authentic material from Switzerland, Genfer See, J.F.Gaudin (W), belongs 

to Juncus bulbosus.] 

Juncus articulatus var. fluitans Wahlenb., Fl. Lapp. 32 (1812). T: [Sweden, Lappland] Lapponia, 

G.Wahlenberg; syn: UPS, n.v. 

Juncus polycephalus D.Don ex Hook., Fl. Scot. 109 (1821), nom. illeg., non Michx. (1803); Juncus 

multicapitatus Schult. & Schult.f., Syst. Veg. 7(1): 198 (1829); Juncus nigritellus D.Don, in G.E.Smith, Engl. 

Bot. Suppl. 1, tab. 2643 (1830), nom. illeg.; Juncus lampocarpus subsp. nigritellus (Macreight) Nyman, 

Consp. Fl. Eur. 748 (1882); Juncus lampocarpus var. nigritellus [D.Don] Macreight, Man. Brit. Bot.: 242 

(1837); Juncus supinus var. nigritellus (Macreight) F.W.Schultz, Flora 23(2): 640 (1840); Juncus 

lampocarpus var. pauciflorus Sond., Fl. Hamburg. 193 (1851); Juncus articulatus subsp. nigritellus (D.Don) 

249

250 


K.Richt., Pl. Eur. 1: 180 (1890); Juncus articulatus f. pauciflorus (Sond.) Sóo, Acta Bot. Acad. Sci. Hung. 16: 

366 (1971). T: [Great Britain, Scotland] Scotch Alps, G.Don; holo: n.v. 

Juncus macrocephalus Viv., Fl. Cors. Prodr. 5 (1824); Juncus lampocarpus var. macrocephalus (Viv.) Döll, 

Rhein. Fl. 179 (1843); Juncus articulatus subsp. macrocephalus (Viv.) K.Richt., Pl. Eur. 1: 180 (1890); 

Juncus lampocarpus subsp. macrocephalus (Viv.) Rouy, Fl. France 13: 240 (1912). T: Corse, Bonifacio, 

D.Viviani; syn: G, n.v. 

Juncus lampocarpus var. suberectus Spenn., Fl. Friburg. 1: 183 (1825). T: not designated. 

Juncus lampocarpus var. alpinus Spenn., Fl. Friburg. 1: 183 (1825). T: in m. Feldberg, Kandel etc.; syn: n.v. 

Juncus tricephalus J.Gay ex Laharpe, Essai Monogr. Joncac. 44 (1825). T: [Italy] Calabre, L.Thomas; syn: 

K, herb. J.Gay, n.v.; [France] Corse, circa Bonifacio, [P.Thomas]; syn: K, herb. J.Gay. 

Juncus lampocarpus γ [unranked] patens Ser. ex Gaudin, Fl. Helv. 2: 552 (1828). T: [Switzerland] 

Gümlingen, N.C.Seringe, Gram. Exs., no 79; syn: n.v. 

Juncus longicapsularis Chevall., Fl. Gén. Env. Paris 2(1): 279 (1827). T: [Paris], F.F.Chevallier; syn: P, n.v. 

Juncus acutiflorus γ. tenuifolius Gaudin, Fl. Helv. 2: 551 (1828). T: [Switzerland] prope Promenthou, 

J.P.Monnard; holo: n.v. 

Juncus lampocarpus B. [unranked] viviparus Gaudin, Fl. Helvet. 2: 252 (1828); Juncus lampocarpus var. 

prolifer Parl., Fl. Ital. 2: 340 (1852); Juncus lampocarpus f. viviparus Neuman, Sver. Fl. 656 (1901). 


Juncus articulatus [Spielart] radicans G.Mey., Chloris Han. 567 (1836). T: [Germany] Blankenburg im 

Braunschweigschen, G.Meyer; syn: n.v. 

Juncus articulatus [Spielart] fluitans G.Mey., Chloris Han. 567 (1836), nom. illeg.; Juncus articulatus var. 

fluitans G.Mey., Fl. Hanov. Excurs. 586 (1849), nom. illeg. T: [Germany] Bremen, G.Meyer; syn: n.v. 

Juncus articulatus [Spielart] maritimus G.Mey., Chloris Han. 567 (1836), nom. inval.; Juncus articulatus var. 

maritimus G.Mey., Fl. Hanov. Excurs. 586 (1849). T: [Germany] Hannover, Nordseeinseln, G.Meyer; syn: n.v. 

Juncus lampocarpus var. fluitans W.D.J.Koch, Syn. Fl. Germ. Helv. 730 (1837). T: not designated. 

Juncus lampocarpus var. macrocarpus Döll, Rhein. Fl. 179 (1843), n.v.; Juncus articulatus f. macrocarpus 

(Döll) Sóo, Acta Bot. Acad. Sci. Hung. 16: 366 (1971). T: [Germany] Rheineck, Custer; syn: n.v. 

Juncus castelli Tineo, in G.Gussone, Fl. Sicul. Syn. 2: 816 (1845). T: [Italy] Messina, V.Tineo; syn: FI, n.v. 

Juncus articulatus var. littoralis Patze, E.Mey. & Elkan, Fl. Prov. Preussen: 67 (1850); Juncus lampocarpus 

var. littoralis (Patze, E.Mey. & Elkan) Buchenau, Bot. Jahrb. Syst. 12: 379 (1890). T: not designated. 

Juncus articulatus var. setiformis Patze, E.Mey. & Elkan, Fl. Prov. Preussen: 67 (1850). T: not designated. 

Juncus lampocarpus [unranked] virescens Döll, Fl. Baden 1: 329 (1857). T: Baden, Neuburgweier, 

J.C.Döll; syn: n.v. 

Juncus lampocarpus var. repens Sond., Fl. Hamburg. 193 (1851); Juncus lampocarpus B. [unranked] repens 

Asch. & Graebn., Fl. Nordostdeutschl. Flachl. 177 (1898). T: [Germany, Hamburg, otherwise not indicated; 

a reference to E.F.Nolte, Nov. Fl. Holsat. (1836)]. 

Juncus gussonii Parl., Fl. Ital. 2: 341 (1852). T: numerous syntypes from Sicily, mainly vicinity of 

Palermo, e.g.: Fiume Grande presso Termini, Cattolica, G.Gussone; syn: FI. 

Juncus lampocarpus var. repens Parl., Fl. Ital. 2: 340 (1852), nom. illeg. T: [Italy] Pisa [collector not 

given]; syn: FI, n.v. 

Juncus lampocarpus var. multiflorus Lange, Haandb. Dansk Fl. 3: 264 (1864), n.v. [fide Lange, in 

M.Willkomm & J.M.C.Lange, Prodr. Fl. Hispan. 1: 184 (1861)]. T: Prope urbem Malaga ad rivum arroyo 

de los Angeles, 20 May 1845, M.Willkomm; syn: COI. 

Juncus lampocarpus b. [unranked] pallidus Schur, Enum. Pl. Transsilv. 685 (1866). T: [Romania] 

Hermannstadt, Kronstadt, F.Schur; syn: n.v. 

Juncus lampocarpus d. [unranked] alpigenus Schur, Enum. Pl. Transsilv. 685 (1866); Juncus articulatus f. alpigenus 

(Schur) Sóo, Acta Bot. Acad. Sci. Hung. 16: 366 (1971). T: [Romania] Arpas, Butian, F.Schur; syn: n.v. 

Juncus lampocarpus e. [unranked] viviparus Schur, Enum. Pl. Transsilv. 685 (1866). T: [Romania] 


Juncus lampocarpus e. [unranked] prolifer Schur, Enum. Pl. Transsilv. 685 (1866). T: [Romania] 


Juncus acutiflorus a. [unranked] pallidiflorus Schur, Enum. Pl. Transsilv. 686 (1866). T: [Romania] 

Hermannstadt, F.Schur; syn: n.v.; [Romania] am Budös bei Tusnad, F.Schur; syn: n.v. 

Juncus lampocarpus var. erectus Čelak., Prodr. Fl. Böhmen. 1: 81 (1867). T: not designated.


Juncus articulatus var. obtusatus Engelm., in A.Gray, Manual, ed. 5, 541 (1867); Engelm., Trans. Acad. Sci. 

St. Louis 2: 497 (1868); Juncus amblyocarpus Rydb., Brittonia 1: 85 (1931). T: [USA] Kaighu's Point, 

Delaware near Philadelphia, Jul 1866, E.Diffenbaugh & E.Burke [G.Engelmann, Herb. Junc. Bor.-Amer. 

Norm.] 50; syn: MO, PR. 

Juncus articulatus var. nigrescens Lindem., Prodr. Fl. Cherson. 206 (1872). T: not designated. 

Juncus articulatus var. pallescens Lindem., Prodr. Fl. Cherson. 206 (1872). T: not designated. 

Juncus articulatus var. subalpinus Borbás, Math. Természettud. Kozlem. 12: 348 (1878). T: [Romania] ad 

rivulos et locis turfosis Zsudjele alpium Retyezat, V.Borbás; syn: BP, n.v.; in valle Groapa Bisztri alpium 

Szarko, V.Borbás; syn: BP, n.v. 

Juncus lampocarpus f. parviflorus Racib., Spraw. Komis. Fizyjogr. 22(2): [p. separ.] 15 (1888); Juncus 

lampocarpus var. parviflorus (Racib.) Zapał., Consp. Fl. Gallic. Crit. 1: 137 (1906). T: [Poland] 

Niepołomice, A.Krupa; syn: n.v.; Korbielów, A.Krupa; syn: n.v. 

Juncus lampocarpus var. cuspidatus Brenner, Meddel. Soc. Fauna Fl. Fenn. 16: 57 (1889); Juncus articulatus 

var. cuspidatus (Brenner) Maire, Fl. Afr. Nord. 4: 302 (1957). T: Finland, Nylandia, Kyrkslätt, Kalliojärvi, 

1886, M.M.W.Brenner; syn: H. 

Juncus lampocarpus f. brachycarpus Trab., in J.A.Battandier & L.C.Trabut, Fl. Algérie, ed. 2, 84 (1895); 

Juncus articulatus var. brachycarpus (Trab.) Maire, Fl. Afr. Nord. 4: 301 (1957). T: Algeria, Cherchell, 

L.C.Trabut; syn: n.v.; [a specimen under this name but without locality or date, signed by Trabut]; syn: P. 

Juncus lampocarpus var. corymbosus Neuman, Sver. Fl. 656 (1901). T: not designated. 

Juncus lampocarpus var. subobtusus Neuman, Sver. Fl. 656 (1901). T: not designated. 

Juncus lampocarpus f. nanus Neuman, Sver. Fl. 656 (1901). T: not designated. 

Juncus lampocarpus [unranked] viridiflorus Asch. & Graebn., Syn. Mitteleur. Fl. 2(2): 478 (1904); Juncus 

articulatus f. viridiflorus I.Grinţ., in E.J.Nyárády, Fl. R. S. 11: 564 (1966). T: not designated. 

Juncus lampocarpus [unranked] pallidiflorus Asch. & Graebn., Syn. Mitteleur. Fl. 2(2): 478 (1904). 


Juncus lampocarpus [unranked] niger Asch. & Graebn., Syn. Mitteleur. Fl. 2(2): 478 (1904). T: not designated. 

Juncus lampocarpus [unranked] pallescens Asch. & Graebn., Syn. Mitteleur. Fl. 2(2): 478 (1904). T: not 

designated. 

Juncus lampocarpus [unranked] congestus Asch. & Graebn., Syn. Mitteleur. Fl. 2(2): 478 (1904). T: not 

designated. 

Juncus lampocarpus [unranked] alpicola Asch. & Graebn., Syn. Mitteleur. Fl. 2(2): 478 (1904); Juncus 

articulatus f. alpicola (Asch. & Graebn.) Sóo, Acta Bot. Acad. Sci. Hung. 16: 366 (1971). T: not designated. 

Juncus lampocarpus [unranked] subobtusatus Asch. & Graebn., Syn. Mitteleur. Fl. 2(2): 479 (1904); Juncus 

articulatus f. subobtusatus (Asch. & Graebn.) Sóo, Acta Bot. Acad. Sci. Hung. 16: 366 (1971). T: not designated. 

Juncus lampocarpus var. senescens Buchenau, Bot. Jahrb. Syst. 37, Suppl. 82: 19 (1905). T: [China, 

N Shaanxi; citation:] T'ai pa shan, Ki fon shan, Quae ssu eel ti u.a.O. [=and other sites], G.Giraldi 6720, 6721, 

6728, 6729, 6734–6738; syn: W, n.v. 

Juncus lampocarpus f. viridiflorus Zapał., Consp. Fl. Gallic. Crit. 1: 137 (1906), nom. inval., non Asch. & 

Graebn. (1904); Juncus articulatus f. zapalowiczii Sóo, Acta Bot. Acad. Sci. Hung. 16: 366 (1971), nom. inval. 

Juncus lampocarpus var. caespitosus Zapał., Consp. Fl. Gallic. Crit. 1: 137 (1906). T: [Poland] Niwra, 

Lenz; holo: n.v. 

Juncus lampocarpus f. nigrescens Zapał., Consp. Fl. Gallic. Crit. 1: 138 (1906); Juncus articulatus f. 

nigrescens (Zapał.) Sóo, Acta Bot. Acad. Sci. Hung. 16: 366 (1971), nom.illeg., non Lindemann (1872). 

T: [Poland] w górach Pokuckich, H.Zapałowicz; syn: n.v. 

Juncus vallis-demonis Lojac., Fl. Sicul. 3: 158 (1909). T: [Italy, Sicily] Fiume di Cannata,Val Demone, Jul 

1853, Reina; holo: PAL. 

Juncus lampocarpus f. submersus Glück, Biol. Morphol. Untersuch. Wasser- Sumpfgew. 3: 219 (1911), n.v.; 

Juncus articulatus f. natans (Glück) Sóo, Acta Bot. Acad. Sci. Hung. 16: 366 (1971). T: n.v. 

Juncus lampocarpus f. natans Glück, Biol. Morphol. Untersuch. Wasser- Sumpfgew. 3: 219 (1911), n.v.; 

Juncus articulatus f. submersus (Glück) Sóo, Acta Bot. Acad. Sci. Hung. 16: 366 (1971). T: n.v. 

Juncus lampocarpus f. maximus Erdner, Ber. Naturwiss. Vereins. Schwaben Neuburg 39–40: 113 (1911). 

T: [Germany, Schwaben, Neuburg a. D.] Ufersand des Lechs [Lech R.] zwischen Feldheim und Marxheim, 

E.Erdner; syn: n.v. 

Juncus lampocarpus var. lindhardii Wiinst., in C.C.Raunkiaer, Dansk Ekskurs.-Fl. 65 (1934); Juncus 

articulatus var. lindhardii (Wiinst.) Wiinst., in C.C.Raunkiaer & K.J.F.Wiinstedt, Dansk Ekskurs.-Fl. 65 

(1942). T: Denmark, Seeland, Kikhavn, 1929, J.Lindhard; lecto: C, designated here by L.Hämet-Ahti. 

251

252 


Juncus subarticulatus Zakirov & Novopokr., Bot. Mater. Gerb. Bot. Inst. Komarova Akad. Nauk SSSR 12: 66 

(1950). T: Zeravschan, lacus Kara-Kul, 13 Aug 1940, K.Z.Zakirov, M.G.Popov & P.Sergeeva; holo: LE. 

Juncus articulatus var. hylandri Hämet-Ahti, Acta Bot. Fenn. 72: 4 (1966). T: Finland, Nylandia, Pyhtää, 

Purola, 1965, A.Saarisalo-Taubert & L.Hämet-Ahti; holo: H; iso: TUR, UPS. 

Juncus articulatus [Spielart] coarctatus G.Mey., Chloris Han. 567 (1836), nom. inval. 

[Juncus repens auct., ascribed to E.F.Nolte, Nov. Fl. Holsat. 38 (1826) - not a name, only a later ascribed 

authorship] 

Illustrations: T.Szynal & J.Mądalski, Atlas Fl. Pol. 2(2): 140 (1931); Ö.Nilsson & S.Snogerup, Bot. Not. 125: 

137, fig. 74 (1972); M.C.Fernández-Carvajal, in B.Valdés et al. (eds), Fl. Vasc. Andalucia Occid. 3: 227 (1987). 

Maps: H.Meusel, E.Jäger & E.Weinert, Vergl. Chorol. Zentraleur. Fl. 1: 85 (1965); E.Hultén & M.Fries, Atlas 

N.Eur. Vasc. Pl. 1: 94, map 188 (1986); E.Hultén, Fl. Alaska 292 (1968). 

Plants mid-green, deep green or paler green; rhizome short, short-creeping, rarely with nodes 

elongated; stems sometimes prostrate, submerged or growing in moving sand and rooting at 

basal nodes. Inflorescence decompound, usually 3–8 × 2–5 cm, with main branches ±erectopatent, 

of (1–) 10–35 (–80) usually obconic (less often semiglobose to subglobose) usually 

(2–) 4–14 (–20)-flowered heads. Tepals equal or subequal, (1.8–) 2.2–3.5 mm long; outer 

tepals sometimes slightly longer, carinate, ±narrowly bordered, varying in colour from green 

dorsally with darker lateral bands and apex dark brown or castaneous-coloured, acute or 

rarely subobtuse and mucronate; inner tepals obtuse to acute, often mucronate, with distinct 

scarious margins. Anthers (0.4–) 0.5–0.9 mm long, usually ±equalling filaments, rarely 

slightly longer or shorter; style 0.2–0.3 mm long; stigmas to 1.5 mm long. Capsule 

trigonous-ovoid, (2.5–) 2.9–4.0 mm long, acute, seldom subobtuse to obtuse, exceeding 

perianth, usually pale brown to castaneous-brown; rostrum c. 0.3 mm long. 2n = 80, fide 

N.A.Harriman & D.Redmond, Rhodora 78: 733 (1976); R.L.Taylor & G.A.Mulligan, Fl. 

Queen Charlotte Is. 2: 43 (1968); M.Zandee, Proc. Kon. Ned. Akad. Wetensch., C, 84: 250 

(1981). Fig. 109. 

A taxon with one of the largest geographical ranges in the genus. Common in most of 

Eurasia and North America, North Africa, and probably introduced in many regions of the 

Southern Hemisphere. 10: DEN, FIN, FOR, GRB, ICE, IRE, NOR, SWE. 11: All. 12: All. 

13: All. 14: All. 20: ALG, LBY, MOR, TUN. 21: AZO, CNY, MDR. 30: ALT, BRY, CTA, 

IRK, KRA, WSB, YAK. 31: AMU, KAM, KHA, KUR, SAK. 32: All. 33: NCS, TCS. 34: 

AFG, CYP, EAI, IRN, IRQ, LBS, PAL, TUR. 36: CHC, CHI, CHM, CHN, CHQ, CHS, CHT, 

CHX. 37: MON. 38: JAP, KOR. 40: EHM, IND, NEP, PAK, WHM. 41: VIE. 50: nsw, vic, 

tas, soa, wau. 51: atp, ctm, nzn, nzs. 70: ASK. 71: BRC. 72: LAB, NBR, NFL, ONT, PEI, 

QUE. 73: COL, IDA, ORE, WAS. 74: MIN, NEB, SDA. 75: CNT, INI, MAI, MAS, MIC, 

NWH, NWJ, NWY, OHI, PEN, RHO, VER, WVA. 76: ARI, NEV, UTA. 77: NWM. 

78: KTY, VRG. 83: per. In a variety of wet habitats, such as pastures and meadows, shores, 

wet tracks, seepage sites, road verges etc. Map 394. 

10. ICELAND: Kjósarsýsla, Úlfarsa, K.Egilsson 144 (K). 11. GERMANY: Oberfranken, Bamberg, 

G.Fischer [Fl. Exs. Bavar.] 578a (PR). POLAND: Szczeczyn, A.Lüderwaldt [A.Kneucker, Cyper. Junc. 

Exs.] 267 (PR, PRA). 12. PORTUGAL: Coimbra, mata do Escarbote, Jun 1892, M.Ferreira (COI). 

SPAIN: Asturias, Luarca, San Cristóbal, 7 Sep 1972, T.E.Díaz (FCO23567); prope urbem Malaga ad rivum 

arroyo de los Angeles, 20 May 1845, H.M.Willkomm (COI). 13. GREECE: Attika, Marathon, Jun 1933, 

L.C.Pinatzi 42 (W). 34. AFGHANISTAN: Salang Pass, 2200 m, K.H.Rechinger 31648 (W). IRAN: 

Mazanderan, Haraz Valley, Panjab, P.Wendelbo 1662 (W). LEBANON: Djebel Sannin, E.Hartmann 

[A.Kneucker, Cyper.Junc. Exs.] 85 (PR, PRA). TURKEY: Ağri, 13 km S of Iğdir, N.Jardine 575 (E, W). 

36. XINJIANG: Chinese Turkestan, Thian Shan, 1130 m, C.Persson 397 (S). CHINA SOUTH-CENTRAL: 

Sichuan, Taofu, Lhamo, Mondeh La, 3700 m, H.Smith 12418 (S, UPS). 

Goulburn, B.Kaspiew 419 (L). 

50. NEW SOUTH WALES: 

Some of the variants of subsp. articulatus are quite distinct, with a well-defined geographical 

range. This particularly applies to "var. hylandri" and "var. lindhardii", both confined to 

N Europe. Even after the exclusion of the following subspecies, subsp. articulatus remains a very 

variable taxon. One of the most conspicuous variants is the large-flowered form from S and 

SW Europe, sometimes recognized as subsp. macrocephalus. Further study is needed. Subspecies 

articulatus is an alloploid sharing genomes with J. acutiflorus and J. alpinoarticulatus, fide 

E.W.Timm & A.R.Clapham, New Phytol. 39: 2–4 (1940), S.Riebe, Bot. Not. 131: 269–272.


Figure 109. Juncus articulatus subsp. articulatus. A–D, habit and inflorescences; E, details 

of inflorescence infested by Livia juncorum; F, tepals and stamens; G, H, perianth and 

capsule. Scale bars: A–E = 3 cm; F = 1 mm; G, H = 2 mm. Drawn by Ö.Nilsson. Reproduced 

with permission, from Ö.Nilsson & S.Snogerup, Bot. Not. 125: 137, fig. 74 (1972). © Lund 

Botanical Society. 

253

254 


192b. Juncus articulatus subsp. limosus (Worosch.) Worosch., in A.K.Skvortsov, 

Florist. Issled. Raz. Raion. SSSR 157 (1985) 

Juncus limosus Worosch., Bjull. Glavn. Bot. Sada 68: 47 (1968). T: Russia, Khabarovskiy kray, Mariinsk, 

12 Aug 1965, V.N.Woroschilow 12475; holo: MHA. 

Juncus lampocarpus var. turczaninowii Buchenau, Bot. Jahrb. Syst. 12: 378 (1890); Juncus turczaninowii 

(Buchenau) Freyn, Oesterr. Bot. Z. 53: 23 (1903); Juncus alpinus subsp. turczaninowii (Buchenau) Worosch., 

in A.K.Skvortsov, Florist. Issled. Raz. Raion. SSSR 157 (1985). T: [Russia, Chita, Nerchinsk] In humidis 

Dahuriae Nerczynensis, 1831, P.K.N.S.Turczaninow; holo: LE. 

Juncus jeholensis Satake, Rep. Sci. Exped. Manchoukuo, sect. 4 [=Index Fl. Jehol.], 4: 106 (1936); Juncus 

turczaninowii var. jeholensis (Satake) K.F.Wu & Ma, in Ma Yu-chuan, Fu Hiang-chian & Chen Shan, Fl. 

Intramongol. 8: 157 (1985); also reportedly in Acta Sci. Nat. Univ. Intramongol. 15(1): 111 (1984), n.v. 

T: Manchuria, Prov. Hsing-an occid., circa Pa-lin-ch’iao, 28 Sep 1933, T.Nakai, M.Honda & M.Kitagawa; holo: TI. 

Illustrations: V.S.Novikov, in S.S.Kharkevich, Sosud. Rast. Soviet. Dal. Vost. 1: 71, tab. 5 (1985); Ma Yuchuan, 

in Ma Yu-chuan, Fu Hiang-chian & Chen Shan, Fl. Intramongol. 8: 158 (1985); Wu Kuofang, Fl. R. P. 

Sinicae 13(3): 183, tab. 42 (1997). 

Maps: H.Meusel, E.Jäger & E.Weinert, Vergl. Chorol. Zentraleur. Fl. 1: 85 (1965); V.S.Novikov, in 

S.S.Kharkevich, Sosud. Rast. Soviet. Dal. Vost. 1: 78, fig. 23 (1985) [the Russian Far East]; N.K.Kovtonyuk, 

in L.I.Malyshev & G.A.Peshkova, Fl. Sibiri 4 (Araceae – Orchidaceae): 163, map 32 (1987) [Siberia]. 

Plants pale green, 20–50 cm tall; rhizome short-creeping. Inflorescence decompound, of 4–25 

usually semiglobose 6–9-flowered heads. Tepals ±equal; outer tepals sometimes slightly 

longer, (1.8–) 2.2–2.8 mm long, carinate, broadly bordered, greenish, with central band often 

bordered brownish near apex, acute; inner tepals (1.8–) 2.4–2.8 mm long, acute to subobtuse 

and mucronate, green, distally broadly bordered scarious. Anthers 0.5–0.7 mm long; 

filaments 1.0–1.1 mm long; style c. 0.2 long; stigmas c. 1.0 mm long. Capsule oblong-ovoid, 

trigonous, exceeding perianth; rostrum 0.3–0.4 (–0.5) mm long. 

Known from the Far East, most common in the Amur and Khabarovsk regions and the 

adjacent regions of Chinese Manchuria. 30: BRY, CTA, YAK. 31: AMU, KAM, KHA, KUR, 

PRM, SAK. 36: CHI, CHM. 37: MON. 38: JAP. Wet meadows, shores of pools. Map 395. 

31. KHABAROVSK: 2 km SE Sofiisk, 1 Aug 1978, A.A.Nechaev s.n. (PRA). 

193. Juncus scheuchzerioides Gaudich., Ann. Sci. Nat. (Paris) 5: 100 (1825) 

T: Iles Malouines, 1820, Gaudichaud; lecto: P, fide A.Lourteig, Publ. Comit. Nat. Franc. Rech. Antarct., Biol. 

23: 41 (1968); iso: K, P, S. 

Juncus inconspicuus d'Urv., Mém. Soc. Linn. Paris 4: 603 (1826); Juncus scheuchzerioides var. inconspicuus 

(d'Urv.) Hook.f., Fl. Antarct. 1: 80 (1844). T: Isla Soledad, D.d’Urville; holo: P; iso: P. 

Illustrations: F.Buchenau, Junc. S.-Amer., tab. 4 (1879); F.Buchenau, in H.G.A.Engler, Pflanzenr. (iv.36) 25: 

170, fig. 86 (1906); M.Barros, Darwiniana 10: 415, fig. 43 (1953); A.Lourteig, Publ. Comit. Nat. Franc. 

Rech. Antarct., Biol. 23: 40, fig. 2 (1968). 

Map: A.Lourteig, Publ. Comit. Nat. Franc. Rech. Antarct., Biol. 23: 38 (1968). 

Perennials, c. 2–20 cm tall, caespitose; rhizome long, creeping-ascending, rooting, distally, 

much branched, wiry, castaneous to pale brown. Stem base covered with stramineous scales. 

Leaves tufted, both basal and cauline, unitubular, terete to ±compressed, perfectly septate, 

acute, c. 3–10 (–20) cm long, 0.4–1.0 mm in diam., exceeding stems; sheaths broad, 

conspicuous, pale green to stramineous, with 2–4 veins on each side; sheath margins broad, 

stramineous to scarious; auricles obtuse, scarious, 0.3–1.0 mm long. Lower bract with a 

broad loose sheath and short acute blade, c. 0.5–7 [in Macquarie Is. specimens] cm long, 

usually exceeding inflorescence. Inflorescence a single head (or with 2 extra heads; or up to 

3 clusters in Macquarie Is. specimens); head sessile, cup-shaped, 3–5 (8)-flowered, to 8–9 mm 

in diam. Tepals ±equal, or outer slightly longer, ±lanceolate, (3.5 in Macquarie Is. 

specimens)–5.2 mm long; outer tepals subacute; inner tepals ±obtuse; central band greenish; 

base distinctly veined; tepal border castaneous to deep brown. Stamens 6, subequalling 

tepals; anthers 0.7–1.9 mm long (0.7 mm in Campbell Is. specimens, 0.8–1.0 mm in 

Macquarie Is. specimens); connective sometimes minutely mucronate (c. 0.2 mm long); 

filaments 1.5–2.5 mm long; style 0.7–1.5 mm long; stigmas 1.5–2.5 mm long. Capsule 

unilocular, oblong-ovoid to ellipsoid-trigonous, ±acuminate, c. 3.5–4.3 mm long, castaneous


to blackish-brown, ±shiny, usually slightly exceeding (rarely equalling) perianth. Seeds ±ovoid, 

c. 0.5–0.6 × 0.3–0.4 mm, reticulate, pale castaneous-brown; appendages absent. 2n = 40, fide 

D.M.Moore, Bot. Not. 113: 187 (1960); D.M.Moore, Bull. Brit. Antarc. Surv. 14: 73 

(1967), n.v. Fig. 110. 

Widely distributed in Subantarctic islands, in southern South America and Antipodes Islands. 

51: ATP. 85: AGS, AGW, CLC, CLN, CLS. 90: CRZ, FAL, KEG, MAQ, MPE, SGE. 

Oligotrophic wet sites, often on peaty or sandy soils from sea level to c. 3000 m. Map 396. 

51. AUCKLAND IS.: Meggs Hill, 28 Dec 1962, E.J.Godley (CHR, P). CAMPBELL IS.: Tucker Cove, 6 Jan 

1961, E.J.Godley (CHR, P); Campbell Island: Davis Point, 16 Jan 1961, V.D.Zotov s.n. (CHR, K, NSW91072). 

85. CHILE SOUTH: Patagonia, Punta Arenas, P.K.H.Dusén 114 (LD); Hermite Island, Cape Horn, J.D.Hooker 

s.n. (NSW254838). 90. FALKLAND IS.: West Falkland, Pt Philomel, Halfway Cove, 21 Nov 1907, 

C.Skottsberg 35 (BA, LD); Stanley Harbour, 30 Mar 1902, C.Skottsberg (LD). KERGUELEN IS.: 

Kerguelens Land, 1839–1843, J.D.Hooker s.n. (NSW254789). MACQUARIE IS.: N end of island, 

T.F.Cheeseman s.n. (NSW254836); Hazelborough Bay, 9 Sep 1950, Taylor s.n. (MEL, NSW254843). 

The material seen from South America and Falkland Is. has stamens with minutely 

mucronate connective, a feature not or seldom observed in plants from the other regions. 

This may indicate a separate status of the South American plants but further study is needed. 

194. Juncus pusillus Buchenau, Junc. S. Amer. 395 (1879) 

Juncus capillaceus Hook.f., Fl. Nov.-Zel. 1: 264 (1853), nom. illeg., non Lam. (1789). T: New Zealand, North 

Island, East Coast, Hawkes Bay, near Eparaima, J.W.Colenso 3825; lecto: K, fide A.Lourteig, Publ. Comit. 

Nat. Franc. Rech. Antarct., Biol. 23: 42 (1968); iso: BM, P, W, WELT. 

Perennials, c. 1–6 cm tall, slender, caespitose to mat-forming; rhizome creeping-ascending, 

distally branched, thin, pale brownish, gradually changing into stem with ascending base. 

Leaves basal and cauline, 2–5, filiform, striate, terete, unitubular, perfectly septate, with 

septa scarcely visible externally, usually exceeding stem, 0.3–0.5 mm in diam.; sheaths 

conspicuous, usually stramineous, loose, with broad membranous or stramineous margins, 

distinctly veined; auricles scarious to stramineous, ±obtuse, c. 0.3–0.5 mm long. Lower bract 

often sheath-like with a short blade or ±leaf-like, 3–7 (–15) mm long, usually exceeding 

inflorescence, often greatly so. Inflorescence terminal, few-flowered, usually of a single 

1–3 (–4)-flowered head. Tepals subequal; inner tepals often longer, often obtuse or slightly 

auriculate, 1.5–2.3 mm long, ±ovate-lanceolate; outer tepals subobtuse, greenish to 

stramineous, later castaneous, distinctly veined; margins membranous. Stamens 6, ±equalling 

tepals; anthers 0.4–0.7 mm long; filaments c. 1.3–1.8 mm long; style 0.3–0.5 mm long; 

stigmas 0.6–1.5 mm long. Capsule unilocular, narrowly ovoid to ellipsoid, subabruptly 

acuminate, to subobtuse, mucronate, c. 1.8–2.5 mm long, brown to castaneous-brown, 

usually slightly longer than perianth. Seeds ±ovoid, c. 0.4–0.5 × 0.25–0.3 mm, pale brown, 

distinctly reticulate; appendages absent. 

Confined to New Zealand, including Stewart Is. and Auckland Is. 51: ATP, NZN, NZS. 

Swampy sites from sea level to 1500 m. Map 397. 

51. AUCKLAND IS.: Ocean Island, Port Ross, Fineran 1933 (CHR, NSW); Hooker Hills, Port Ross, Fineran 

2135 (CHR, NSW). NORTH IS.: Mt Ruapehu, Jan 1875, S.Berggren (LD); Mt Egmont, 3 Jan 1963, B.Allen 

s.n. (CHR, K, NSW91057, P); Makirikiri [c. 160 km N of Wellington], Ruahine Mountains, North Island, 31 

Dec 1943, V.D.Zotov s.n. (CHR, NSW91058). SOUTH IS.: Otira R., Feb 1874, S.Berggren (LD); George 

Sound, Lake Katherine, 17 Apr 1949, R.Mason 65446 (P); Hooker Valley, c. 8 miles [13 km] SSW of Mt 

Cook, 15 Feb 1966, B.G.Briggs s.n. (CHR, NSW90800); Lake Camp, South Ashburton Valley, Canterbury, 

R.Mason 9420 (CHR, NSW); Sabine River, Nelson Lakes National Park, M.J.Simpson 2548 (CHR, K, NSW). 

Records of J. pusillus from Kerguelen Is. and South America belong to J. scheuchzerioides. 

Tasmanian plants traditionally referred to as J. pusillus were recognized as a separate species, 

J. sandwithii by A.Lourteig, Publ. Comit. Nat. Franc. Rech. Antarct., Biol. 23: 44 (1968). 

255

256 


Figure 110. Juncus scheuchzerioides. A–C, habit; D, inflorescence; E, lower bract; F, flower 

bract; G, outer tepal; H, inner tepal; I, pistil with stamen; J, pistil; K, capsule; L, perianth 

with ripe capsule (A–L, J.D.Hooker, P). Scale bars: A–C = 2.5 cm; D–I, K, L = 2 mm; 

J = 1 mm. Reproduced with permission, from A.Lourteig, Publ. Comit. Nat. Franc. Rech. 

Antarct., Biol. 23: 40, fig. 2 (1968).


195. Juncus stipulatus Nees & Meyen, in F.J.F.Meyen, Observ. Bot. 126, 1843. 

T: Chile, Valparaiso, F.J.F.Meyen s.n.; holo: 'herb. Lucaeno', n.v. [the herbarium not identified; A.Lourteig, 

Publ. Comit. Nat. Franc. Rech. Antarct., Biol. 23: 44 (1968), cited a type specimen as 'Meyer II 1831 Type 

PR' but the specimen was not located at PR recently.] 

Juncus oliganthus Phil., Linnaea 29: 75 (1857). T: Chile, Valdivia, Corral, R.A.Philippi s.n.; holo: SGO; 

iso: BREM, G, K, MO, P. 

Juncus mandonii Buchenau, Abh. Naturwiss. Vereine Bremen 4: 121 (1874). T: Bolivia, La Paz, Larecaja, 

2600–4100 m, G.Mandon 1440; lecto: GOET, fide H.Balslev, Fl. Neotrop. Monogr. 68: 127 (1996); iso: BM, 

BR, G, GH, K, LAU, MO, NY, P, S. 

Illustration: F.Buchenau, Abh. Naturwiss. Vereine Bremen 4: tab. 3, fig. 1–7 (1874). 

Map: H.Balslev, Fl. Neotrop. Monogr. 68: 128, fig. 63 (1996) [Neotropical region]. 

Perennials, c. 2–10 (–20) cm tall, loosely caespitose; rhizome creeping, rooting, distally 

ascending and branched, gradually changing into stem, covered with scarious to stramineous 

scales to 1 cm long. Leaves mostly clustered along the transition zone between rhizome and 

stem, c. 1–8 cm long, ±terete, unitubular, perfectly septate; sheaths 2–15 cm long, with 

scarious margins; auricles obtuse, 0.2–0.8 mm long. Lower bract leaf-like, to 2 cm long; 

sheath slightly shorter than inflorescence; blade acicular. Inflorescence of 1 (rarely 2) 

terminal heads, each head (1–) 2–5-flowered. Tepals ±equal, 2.0–3.5 mm long, lanceolate, 

acute, bordered castaneous or stramineous; margins later scarious; central band green, later 

castaneous. Stamens 6, shorter than tepals; anthers 0.8–1.3 mm long; filaments 1.5–2.4 mm 

long. Capsule unilocular, broadly ellipsoid to ±ovoid, 2.5–3.5 mm long, light brown to dark 

blackish brown, slightly exceeding perianth. Seeds ±ovoid, 0.4–0.6 × c. 0.3 mm, reticulate, 

brown; appendages absent. 

Widely distributed in W South America, from mountains of Colombia to Bolivia and to 

southern Chile and Argentina. Edges of streams, lake shores, boggy habitats and open 

ground. Two varieties are recognized but further study is needed. 

Style 0.4–0.5 mm long. Capsule subabruptly acuminate, distinctly mucronate 195a. var. stipulatus 

Style 0.1–0.3 mm long. Capsule subobtuse, shortly mucronate 195b. var. chilensis 

195a. Juncus stipulatus Nees & Meyen var. stipulatus 

Juncus microcephalus var. pusillus E.Mey., in C.Presl, Reliq. Haenk. 1: 142 (1827). T: Peru, T.Haenke s.n.; 

holo: PR, n.v., perhaps lost. 

Juncus corralensis Phil., Linnaea 33: 269 (1864); Juncus stipulatus var. corralensis (Phil.) Buchenau, Junc. S. 

Amer. 394 (1879). T: Chile, Valdivia, Corral, 1859, Krause s.n.; holo: SGO; iso: SGO, P. 

Illustrations: M.Barros, Darwiniana 10: 360, fig. 26 (1953); M.Barros, Darwiniana 10: 415, fig. 43 (1953); 

A.Lourteig, Publ. Comit. Nat. Franc. Rech. Antarct., Biol. 23: 43, fig. 3 (1968); H.Balslev, Fl. Neotrop. 

Monogr. 68: 125, fig. 61A–C (1996). 


Style 0.4–0.5 (–0.7) mm long. Capsule subabruptly acuminate, distinctly mucronate. Seeds 

with scattered tubercules in the reticulation fields. Fig. 111. 

In the Andes from Colombia, Bolivia, Ecuador and Peru to southern Argentina and Chile. 

83: BOL, CLM, ECU, PER. 85: AGS, AGW, CLN, CLC, CLS. Map 398. 

83. COLOMBIA: Caldas, Páramo del Quindio, 3700–4200 m, F.W.Pennell & T.E.Hazen 9963 (GH, K, NY, 

PH, US). ECUADOR: Imbabura, Laguna Mojanda, 3600–3700 m, H.Balslev & F.Quintana 24122 (AAU, F, 

K, NY, QCA). 85. ARGENTINA NORTHWEST: Río Negro, San Carlos de Bariloche, 770 m, 11 Feb 1905, 

O.Buchtien (L, PR, S, SI). CHILE CENTRAL: Aconcagua, Riecillo, 3000 m, H.Gunckel 3873 (L); Curico, 

Cordillera Peteroa-Planchon, Cañon Rio Claro, 1700 m, Y.Mexia 7884 (K). 

195b. Juncus stipulatus var. chilensis (Gay) Kirschner, in S.Snogerup, P.F.Zika & 

J.Kirschner, Preslia 74: 251 (2002). 

Juncus chilensis Gay, Fl. Chil. 6: 146 (1854). T: Chile, Colchagua, Cordillera de Talcaregüe, S. Fernando, 

1833, C.Gay s.n.; holo: P; iso: BREM, MO, P, SGO. 

257

258 


Figure 111. Juncus stipulatus var. stipulatus. A, habit; B, inflorescence; C, auricles; 

D, tepals; E, perianth with capsule; F, pistil with stamen; G, H, capsule (A, C, H, Meyen, PR 

[nowadays not located]; B, D–G, O.Buchtien, S). Juncus sandwithii. I, habit; J, auricles; 

K, inflorescence; L, lower bract; M, flower bract; N, tepals; O, pistil with stamen; 

P, perianth with capsule; Q, R, capsule (I–R, R.C.Gunn 1414, K, P). Scale bars: A, I = 2.5 cm; 

B, C = 1.8 mm; D–H, J–R = 2 mm. Reproduced with permission, from A.Lourteig, Publ. 

Comit. Nat. Franc. Rech. Antarct., Biol. 23: 43, fig. 3 (1968).


Juncus gayanus Steud., Syn. Pl. Glumac. 2: 300 (1855). T: Chile, Colchagua, Cordillera de Talcaregüe, 

1833, C.Gay s.n.; holo: P; iso: MO. 

Juncus depauperatus Phil., Fl. Atacam. 53 (1860), nom. illeg., non Ten. (1830–1832). T: Antofagasta, 

Atacama Desert, Cachinal de la Sierra, 2100 m, R.A.Philippi s.n.; lecto: SGO, fide H.Balslev, Fl. Neotrop. 

Monogr. 68: 127 (1996); iso: SGO, K. 

Illustrations: F.Buchenau, Junc. S.-Amer., tab. 4 (1879); A.Lourteig, Publ. Comit. Nat. Franc. Rech. Antarct., 

Biol. 23: 46, fig. 4 (1968). 


Style 0.1–0.3 mm long. Capsule subobtuse, shortly mucronate. Seeds with lines of fine 

tubercules across the reticulation fields. Fig. 112. 

Known from Chile and adjacent Argentina, also reported from Bolivia and Peru, but not seen 

during this study. 83: BOL, PER. 85: AGS, AGW, CLC, CLN, CLS. Map 399. 

85. ARGENTINA NORTHWEST: Jujuy, Moreno, 3500 m, R.E.Fries 949 (P, S). CHILE CENTRAL: San 

Fernando, Feb 1831, C.Gay (P). CHILE SOUTH: Magallanes, R. San Martín, 6 Feb 1896, P.K.H.Dusén (UPS). 

196. Juncus sandwithii Lourteig, Publ. Comit. Nat. Franc. Rech. Antarct., Biol. 

23: 44 (1968) 

T: Tasmania, Arthur’s Lake, 17 Jan 1845, R.C.Gunn 1414; holo: P; iso: BM, C, HO, K, P, S, WU [the 

holotype collection in P lacks locality details and the collection number]. 

Illustrations: A.Lourteig, Publ. Comit. Nat. Franc. Rech. Antarct., Biol. 23: 43, fig. 3 (1968); K.L.Wilson & 

L.A.S.Johnson, Telopea 9: 359, fig. 1 (2001). 


Perennials, c. 2–10 cm tall, slender, loosely caespitose; rhizome thin, c. 0.2–0.3 mm in 

diam., creeping-ascending, long-noded, gradually changing into stem, with distal ascending 

extravaginal shoots. Stem ascending, filiform. Leaves setaceous, numerous, ±evenly 

distributed along the most distal part of rhizome and lower 3/4 of stem, ±exceeding stem, 

terete, unitubular, perfectly septate with septa ±visible externally, 0.2–0.7 mm in diam., to 

16 cm long; sheaths ±tight; auricles oblong, obtuse, 0.5–0.8 mm long. Lower bract setaceous, 

0.4–5 (–15) cm long, usually equalling to much exceeding inflorescence. Inflorescence fewflowered, 

of 1–4 (–6) obconical (1–) 2–6 (–10)-flowered heads 2–4 mm wide, usually one 

sessile and one on thin peduncle to 3.5 cm long. Capitulum bracts 2 (–3), ovate-lanceolate, 

acute, membranous to pale stramineous, to c. 2 mm long. Tepals ±equal, linear-lanceolate, 

(1.7–) 2.0–3.3 mm long with distinct membranous margins; outer tepals acuminate; inner 

tepals with subobtuse tip, sometimes cucullate, 3-veined proximally, green to pale green, 

with apex and margins often ±reddish. Stamens 6, shorter than tepals or slightly longer; 

anthers 0.4–0.7 mm long; filaments c. 1.0–1.5 mm long; style c. 0.2–0.7 mm long; stigmas 

1.0–2.5 mm long. Capsule narrowly trigonous-ellipsoid (subprismatic) to narrow-ovoid, 

±unilocular, c. 2.8–4.3 mm long, gradually narrowing in a long rostrum, pale castaneous, 

usually much longer than perianth. Seeds ±ellipsoid, 0.4 × 0.25 mm, apiculate; inner seed 

coat regularly reticulate, with surface of fields minutely tuberculate; outer seed coat tight, 

transversally reticulate; appendages absent. Fig. 111. 

Known from Australia (Tasmania, Victoria and New South Wales), once found in Papua- 

New Guinea. 42: NWG. 50: NSW, TAS, VIC. Wet mountain short-grass sites. Map 400. 

42. NEW GUINEA: Mt Victoria, 2700 m, J.R.Croft (A, BISH, BRI, CANB, E, K, L, LAE61840, M, NSW, 

US). 50. NEW SOUTH WALES: Prussian Creek. Kosciusko National Park, 24 Jan 1976, Thompson 2455 

p.p. (AD, BRI, CBG, CHR, HO, K, L, MEL, MO, NSW); NE edge of Bald Rock dome, Bald Rock National 

Park, K.L.Wilson 6962 (BRI, HO, NSW). TASMANIA: Coal R., A.F.Oldfield 131 (K); Central Plateau, 

Lake St. Clair, c. 800 m, Gibbs 6672 (BM, K); Scotts Peak Road E of Lake Pedder, 20 Feb 1970, W.Curtis s.n. 

(HO, NSW276213); Douglas Creek, just N of Arm River Track crossing, Cradle Mtn -Lake St Clair N.P, 

K.L.Wilson 830a (HO, MEL, MO, NSW). VICTORIA: Jimmys Creek, Grampians road, A.C.Beauglehole 

16432 (MEL, NSW); Lake Catani, Mt Buffalo National Park, E.M.Canning 1789 (CANB, NSW); Upper 

Delegate River, near Bidwell, R.Melville 2955, N.A.Wakefield & Hunter (K, NSW). 

259

260 


Figure 112. Juncus stipulatus var. chilensis. A, habit; B, leaf sheath, auricles and blade; 

C, inflorescence; D, details of inflorescence; E, middle bract; F, flower bract; G, Flower; 

H, tepals; I, pistil with stamen; J, capsule; K, perianth with ripe capsule (A–K, C.Gay, P). 

Juncus novae-zelandiae. L, habit; M, leaf with sheath and auricles; N, middle and flower 

bracts; O, flower; P, outer tepal; Q, inner tepal; R, pistil with stamen; S, capsule; T, perianth 

with capsule (I–T, H.Filhol, P). Scale bars: A, L = 2.5 cm; C = 0.5 cm; B, D–K, M–T = 2 mm. 

Reproduced with permission, from A.Lourteig, Publ. Comit. Nat. Franc. Rech. Antarct., 

Biol. 23: 46, fig. 4 (1968).


197. Juncus novae-zelandiae Hook.f., Fl. New Zealand 1: 264 (1853) 

T: New Zealand, banks of the River Waipoua, 1849, J.W.Colenso 1093; lecto: K, fide E.Edgar, in L.B.Moore 

& E.Edgar, Fl. New Zealand 2: 66 (1970); iso: P, WELT. 

Illustration: A.Lourteig, Publ. Comit. Nat. Franc. Rech. Antarct., Biol. 23: 46, fig. 4 (1968). 


Perennials, 2–10 (–15) cm tall, caespitose to loosely caespitose; rhizome long-noded, 

creeping, distally ascending and branched; stem to c. 0.4 mm in diam., often exceeded by 

leaves. Leaves numerous, both basal and inserted along stem; blade unitubular, subterete 

(±compressed), perfectly septate, to c. 5–8 cm long, 0.3–0.6 (–0.9) mm wide; sheaths loose, 

with broad membranous margins, distally veined; auricles obtuse, ±stramineous, 0.3–0.8 mm 

long. Lower bract leaf-like, filiform, ±equalling inflorescence or longer, 0.3–1.3 cm long. 

Inflorescence a single terminal 2–5-flowered head 5–6 mm wide, or the sessile head with 1–3 

shortly pedunculate 1–3-flowered loose clusters. Capitulum bracts broadly ovate, acute to 

obtuse, stramineous to pale castaneous, c. 1.5–2.5 mm long. Tepals subequal; inner tepals 

often slightly longer, (1.5–) 1.9–2.5 mm long, ±obtuse, castaneous, with margins very broad 

and usually scarious to membranous; outer tepals acute, occasionally slightly cucullate and 

carinate, with central band greenish or castaneous, and bordered deep castaneous, with 

margins usually broad. Stamens 6, ±equalling or ±longer than tepals; anthers 0.5–1.0 mm 

long, slightly shorter than c. 0.8–1.1 mm long filaments; style 0.1–0.4 mm long (including 

mucro top); stigmas 1–2 mm long. Capsule broadly ovoid to subglobose when fully ripe, or 

±oblong-ovoid, mucronulate, unilocular, 2.5–3.2 mm long incl. 0.1–0.3 mm mucro, blackish 

castaneous-brown, shiny, conspicuously exceeding perianth. Seeds obliquely ovoid, 

apiculate, 0.4–0.6 × 0.3 mm, reticulate; appendages absent. Fig. 112. 

Endemic to New Zealand. 51: NZN, NZS. Mountain swamps up to 1500 m; less often in 

damp sites in lowlands. Map 401. 

51. NORTH IS.: Wellington, Ruakokopatuna Valley, c. 1000 m, A.J.Healy 50/24 (CHR, K, NSW, P); Mt 

Pureora, R.O.Gardner 971 (AKU, NSW); below Mt Holdsworth, Mangatarereatiwhakatu stream above the 

Donell flat, Tararua Range, R.Melville 5560 & A.P.Druce (K, NSW). SOUTH IS.: Kaimanawa, Feb 1875, 

S.Berggren (LD); Cascade Creek (at South end of Lake Gunn), 25 Feb 1966, B.G.Briggs s.n. (CHR, 

NSW90840); Lake Rotoiti, Nelson, 17 Apr 1965, A.P.Druce s.n. (CHR, NSW 91247); Upper Bealey River, 

Arthurs Pass, Canterbury, L.A.S.Johnson NZ7 (NSW); The Bluff, T.Kirk s.n. (NSW91070); Whites Creek, near 

Mt Oxford, Canterbury, R.Mason 3252 (CHR, K, NSW); Westland, valley below Franz Joseph Glacier, 

R.Melville 6863 & E.F.Melville (K, NSW). 

198. Juncus curtisiae L.A.S.Johnson, in M.R.Banks & al., Asp. Tasman. Bot. Trib. W. 

Curtis 44 (1991) 

T: Tasmania, Table Mountain, 2500 ft, 15 Jan 1975, D.A.Ratkowsky & A.V.Ratkowsky JS101; holo: NSW; iso: 

CANB, HO, MEL. 

Map: L.A.S.Johnson, in M.R.Banks & al., Asp. Tasman. Bot. Trib. W. Curtis 44 (1991). 

Perennials, c. 1–5 cm tall, dwarf, mat-forming; rhizome creeping-ascending, thin, longnoded, 

c. 0.3–0.4 mm in diam., distally branched, with 3 to several leaves on each branch. 

Leaves often densely crowded, to 5 cm long, 0.2–0.5 mm in diam., unitubular, ±terete, 

perfectly septate, ±equalling stem; sheaths ±loose, 3–4-veined on each side, broadly bordered 

membranous; auricles broad, obtuse, c. 0.3–0.4 mm long. Lower bract leaf-like, 0.4–1.5 cm long, 

with loose sheath, ±exceeding inflorescence. Inflorescence of 1 or rarely 2 heads, usually 

2–4-flowered. Capitulum bracts broadly ovate, acute, membranous to scarious, to c. 2 mm 

long. Tepals ±unequal; outer tepals shorter, broadly lanceolate, 1.9–3.0 (–3.7) mm long; 

inner tepals ovate-lanceolate, acute to ±obtuse, longer than or equalling the outer ones, 

reddish castaneous with darker tip, broadly bordered membranous, with central band 

greenish to brownish. Stamens 6, often longer than tepals; anthers 0.6–1.0 (–1.5) mm long; 

filaments c. 1.5–2.2 mm long; style difficult to distinguish from capsule top, c. 0.5–0.7 mm 

long; stigmas 0.7–1.0 mm long. Capsule broadly ellipsoid to ovoid, trigonous distally, 

subabruptly narrowing into a subconical 0.5–1.0 mm rostrum, 2.5–3.8 mm long, golden 

brown to castaneous, slightly exceeding perianth. Seeds ellipsoid-ovoid, apiculate, 0.4–0.6 × 

0.25 mm, regularly reticulate, transversally lineolate; appendages absent. 

261

262 


Endemic to Tasmania. 50: TAS. Boggy sites to c. 1250 m. Map 402. 

50. TASMANIA: Cradle Mtn, Lake Dove, 1240 m, 25 Feb 1970, P.H.Raven 25993 (CHR, MO, NSW, P); Mt 

Wellington, 5 Jan 1975, D.A.Ratkowsky & A.V.Ratkowsky (CHR, HO, NSW); Douglas Creek, just N of Arm 

River Track crossing, Cradle Mtn -Lake St Clair N.P, K.L.Wilson 8308b (CHR, HO, K, NSW); Chimney Pot 

Hill, Mt Wellington, D.A.Ratkowsky JS102 (CHR, NSW); fire road near Trestle Mountain, Mount Wellington, 

D.A.Ratkowsky JS107 & A.V.Ratkowsky (CANB, HO, K, MEL, NSW). 

Very close to J. sandwithii and J. thompsonianus (its mainland Australian counterpart) but 

having reddish-castaneous, broader tepals, inner tepals slightly longer, and a more abruptly 

narrowed capsule. The nature of the differences exclude the possibility that the two taxa are 

conspecific. Collections from Ocean Beach in western Tasmania are much bigger in their 

parts than others (tepals c. 3.7 mm long, for example) and need further study. 

199. Juncus ratkowskyanus L.A.S.Johnson, in M.R.Banks & al., Asp. Tasman. Bot. 

Trib. W. Curtis 43 (1991) 

T: Tasmania, Mount Field National Park, Lake Seal, 27 Jan 1960, R.Carolin 1594; holo: NSW; iso: CANB, 

HO, MEL, SYD. 


Perennials, to c. 2 cm tall, mat-forming; rhizomes thin, later wiry, long-noded, creeping, 

distally branched, ascending. Stems to 0.5 mm in diam., terete, erect. Leaves several, to 2 cm 

long, 0.2–0.6 mm in diam., unitubular, perfectly septate, often exceeding inflorescence; 

sheaths with 1–2 veins on each side; auricles obtuse, 0.5–0.7 mm long. Lower bract shorter 

than to exceeding inflorescence, to 1 cm long; base broadened. Inflorescence a single 

1–3 (–6)-flowered head (very rarely 2 heads). Outer tepals (2.2–) 2.5–3.0 mm long; inner 

tepals slightly longer, all narrowly lanceolate, acute, green or stramineous, sometimes with 

reddish tinge; margins narrowly membranous. Stamens 3; anthers (0.5–) 0.7–1.0 mm long; 

filaments to c. 2 mm long; style 0.5–1.0 mm long; stigmas 0.7–1.0 mm long. Capsule ovoid, 

±abruptly acuminate, dark golden brown to dark castaneous at maturity, equalling or 

occasionally longer than perianth. Seeds ovoid, apiculate, 0.7–1 mm long, reticulate; 


Restricted to Tasmania. 50: TAS. Higher altitude boggy sites. Map 403. 

50. TASMANIA: Mt Wellington, J.H.Hemsley 6710 (NSW); Mt Wellington, Dead Island Bogs, 5 Jan 1975, 

A.V.Ratkowsky (K, NSW); Hartz Mountain National Park, D.A.Ratkowsky & A.V.Ratkowsky JS113 (HO, 

NSW); Lake Seal, Mt Field National Park, R.Carolin 1594 (CANB, HO, MEL, NSW); S end of Thark Ridge, 

Mt Wellington, A.V.Ratkowsky JS105 & D.A.Ratkowsky (CHR, NSW); Dead Island Bogs, Mt Wellington, 

A.V.Ratkowsky JS109 (K, NSW). 

Similar to J. sandwithii but having three stamens, a very low growth, ±abruptly acuminate 

and dark castaneous-brown capsule. 

200. Juncus thompsonianus L.A.S.Johnson, in M.R.Banks & al., Asp. Tasman. Bot. 

Trib. W. Curtis 45 (1991) 

T: New South Wales, Kosciusko National Park, Wragges Creek, 1600 m, 3 Feb 1979, K.L.Wilson 2033; holo: 

NSW; iso: CANB, HO, MEL. 

Illustration: K.L.Wilson, in G.J.Harden (ed.), Fl. New South Wales 4: 288 (1993). 


Perennials, 0.5–3 cm tall, mat-forming; rhizome creeping, slender, long-noded, distally 

branched, ascending and leafy. Stems erect, terete, to c. 0.5 mm in diam. Leaves mostly 

caespitose and also cauline, unitubular, perfectly septate, terete, to 7 cm long, 0.3–0.5 mm in 

diam., usually much exceeding inflorescence; sheaths loose, with broad scarious margins and 

3–4 veins on each side; auricles ±acute, c. 0.5 mm long. Lower bract leaf-like, often 

exceeding inflorescence, to 2 cm long, with base broad, scarious. Inflorescence of a single 

1–3-flowered head (rarely heads 2). Tepals subequal; outer tepals ±ovate, obtuse to acute, 

1.8–2.5 mm long, slightly shorter than narrowly lanceolate; inner tepals broad-acute to 

obtuse, green or stramineous, sometimes with a reddish tinge; margins membranous, broad.


Stamens 6; anthers 0.4–0.8 mm long; filaments c. 1.5 mm long; style 0.3–0.7 mm long; 

stigmas 0.5–1.3 mm long. Capsule ±ovoid, ±abruptly acuminate, c. 2 mm long, pale golden 

brown, slightly exceeding or equalling perianth. Seeds ellipsoid, minutely apiculate, 0.4–0.7 mm 

long, reticulate; appendages absent. 

Confined to New South Wales and Victoria, Australia. 50: NSW, VIC. High altitude 

mountain bogs. Map 404. 

50. NEW SOUTH WALES: Southern Tablelands, Kiandra, Cave Ck, A.Rodd & R.Coveny 2675 (CANB, NSW, 

P); Kosciusko National Park, Perisher Ck, J.Thompson 2699 (NSW, PR); Lower Spencers Creek, Kosciusko 

National Park, J.Thompson 1690 (CHR, HO, NSW); Schlink Pass, Munyang R. at Whites River Hut, 

K.L.Wilson 6147 (NSW, PRA); Thredbo R. near Dead Horse Gap, Kosciusko National Park, J.Thompson 4215 

(AD, CANB, K, MEL, MO, NSW); Wragges Ck, Kosciusko National Park, K.L.Wilson 2033 (CANB, HO, 

MEL, NSW). VICTORIA: Mt Fainter High Plains, Feb 1928, A.J.Tadgell (MEL1533630). 

Presumed Hybrids 

Juncus ×fulvescens Fernald, Rhodora 35: 236 (1933) = Juncus articulatus L. × Juncus 

brevicaudatus (Engelm.) Fernald 

T: USA, Maine, Washington Co., Pembroke, Charlotte Road, 18 Aug 1909, M.L.Fernald 1588; holo: GH. 

Juncus ×alpiniformis Fernald, Rhodora 35: 235 (1933) = Juncus alpinoarticulatus Chaix 

× Juncus articulatus L. 

T: Canada, W Newfoundland, Bonne Bay, Storehouse Cove, 9 Aug 1929, M.L.Fernald, B.Long & J.M.Fogg 

Jr. 1501; holo: GH; iso: K. 

Juncus ×buchenaui Dörfl., Herb. Norm., no. 3267 (1897), and Dörfler, Herb. Norm. Sched. Cent. 33: 95, no. 

3267 (1897), nom. illeg., non Sved. (1897). T: Sweden, Skåne, in litore lacus Ringsjön, Aug. 1894, 

R.Herlitz [I.Dörfler, Herb. Norm.] 3267; syn: PR, W. 

Juncus alpinus × lampocarpus B. [unranked] roeperi Asch. & Graebn., Syn. Mitteleur. Fl. 2 (2): 482 (1904), 

nom. inval. Authentic material: [Germany] Mecklenburg, Dolgen, J.A.C.Roeper; n.v. 

Illustration: Ö.Nilsson & S.Snogerup, Bot. Not. 125: 207, fig. 78 (1972) [J. alpinoarticulatus subsp. 

alpinoarticulatus × J. articulatus subsp. articulatus] & 125: 208, fig. 79 (1972) [J. alpinoarticulatus subsp. 

rariflorus × J. articulatus subsp. articulatus]. 

At least three subspecies of Juncus alpinoarticulatus (subsp. alpinoarticulatus, subsp. 

nodulosus and subsp. americanus) hybridize with Juncus articulatus. The hybrids are seedsterile 

or almost completely so, and they propagate vegetatively. The hybrid proved to have 

2n = 60, fide S.Snogerup, in Ö.Nilsson & S.Snogerup, Bot. Not. 125: 206–207 (1972); 

M.Zandee, Proc. Kon. Ned. Akad. Wetensch., C, 84: 250 (1981). It is known from Europe, 

Turkey and E North America. 10. FIN, GRB, NOR, SWE. 11. AUT, CZE, GER, POL, SWI, 

12. FRA. 13. GRC, ITA, 14. RUS. 34. TUR. 

11. SWITZERLAND: Bern, Beatenberg, 26 Aug 1911, A.Thellung (K). 

Juncus articulatus L. × Juncus nodosus L. 

Reported to occur in North America by R.E.Brooks & S.E.Clemants, Fl. N. Amer. 22: 255 

(2000) without further details; material not seen for this study. 

Juncus ×lemieuxii Boivin, Naturaliste Canad. 94: 525 (1967) = Juncus articulatus L. 

× Juncus canadensis J.Gay ex Laharpe 

T: [Québec] Vaudreuil-Soulanges, Pointe Cascade, le long du Saint-Laurent face à l'Ile Ronde, 9 Aug 1962, 

G.Lemieux 3681; holo: DAO. 

Described from Canada; reported to occur in North America by R.E.Brooks & S.E.Clemants, 

Fl. N. Amer. 22: 255 (2000) without further details; material not seen for this study. 

263

264 


Juncus ×langii Erdner, Allg. Bot. Z. Syst. 12: 196 (1906) = Juncus acutiflorus Ehrh. ex Hoffm. 

× Juncus alpinoarticulatus Chaix 

T: [Germany, Bavaria] Schwaben, Monheim, Jul 1905, E.Erdner; holo: n.v. 

Rare or overlooked hybrid seen and reported from Austria, France, Italy, Poland, 

Switzerland, The Netherlands and Germany. It is seed-sterile, with a great ability to form 

creeping rhizomes. 11. GER, AUT, NET, POL, SWI. 12: FRA 13: ITA. 

11. GERMANY: Bayern, Monheim, 13 Aug 1907, E.Erdner (K, PRA). 

Juncus ×gracilescens F.J.Herm. ex Wadmond, Rhodora 49: 120 (1947) = Juncus 

alpinoarticulatus Chaix × Juncus brevicaudatus (Engelm.) Fernald 

T: New Brunswick, Bathurst, S.F.Blake; syn: n.v.; Ontario, Northern Lake Huron, Whitefish Bay, 30 Aug 

1932, W.Koelz 4681, 4687; syn: MICH. 

Reported from Ontario, New Brunswick and Minnesota. 

Juncus ×nodosiformis Fernald, Rhodora 35: 235 (1933) = Juncus alpinoarticulatus Chaix 

× Juncus nodosus L. 

T: Canada, Québec, St. John River, 23 Aug 1904, J.F.Collins, M.L.Fernald & A.S.Pease; holo: GH. 

A sterile hybrid known from the type collection and from Newfoundland. 

Juncus ×stuckeyi Reinking, Brittonia 33: 175 (1981) = Juncus alpinoarticulatus Chaix 

× Juncus torreyi Coville 

T: USA, Ohio, Erie Co., Lake Erie, Kelleys Is., Deep Northwest Quarry, 29 Jun 1979, M.F.Reinking 612; holo: OS. 

Sterile and intermediate between the parents. Further records come from Erie Co. and Ottawa 

Co., Ohio (deposited in OS). 

Juncus articulatus L × Juncus bulbosus L. 


A completely seed-sterile hybrid, capsules imperfectly developed, vegatatively propagating 

locally. Known from a few places in NE Skåne and S Halland, Sweden. 2n = 60, fide 

S.Snogerup, Bot. Not. 113: 246 (1960). Fig. 113. 

Juncus ×montserratensis Marcet, Bol. Soc. Esp. Hist. Nat. 45: 238 (1947) = Juncus 

acutiflorus Ehrh. ex Hoffm. × Juncus articulatus L. 

Juncus lampocarpus f. acutipetalus Marcet, Bol. Soc. Esp. Hist. Nat. 43: 39 (1945). T: [Spain] Montserrat, 

Fuente de las "Guineus", F.Marcet; holo: BC, n.v. 

Juncus ×surrejanus Druce ex Stace & Lambinon, Bull. Jard. Bot. État. 53: 508 (1983). T: Hedge Court 

Millpond, Surrey, 9 Sep 1893, W.H.Beeby; holo: OXF. 

Juncus micranthus Desv., Obs. Pl. Angers 82 (1818), p.p., nom. rejic. propos. T: [France, Angers] 

N.A.Desvaux; syn: G-DEL. [The other syntypes not seen and probably belonging to J. acutiflorus, as also later 

authentic material at K.] 


Rarely between parents. Seed sterile and vegetatively propagating. Known from Denmark, 

France, Spain, Germany and Great Britain. 2n = 60, fide E.W.Timm & A.R.Clapham, New 

Phytol. 39: 2–4 (1940); M.Zandee, Proc. Kon. Akad. Wet., ser. C, 84: 250 (1981). Fig. 114. 

Juncus ×murbeckii Sagorski, Mitt. Thüring. Bot. Vereins, ser. nov., 16: 37 (1902). [and 

G.Beck, Glasn. Zemaljsk. Muz. Bosni. Hercegovini 15: 198 (1903); Fl. Bosne Herceg. 1: 62 

(1903)] = Juncus anceps Laharpe × Juncus articulatus L. 

T: Herzegovina, Nevesinjsko polje, Žiljevo, 29 Jul 1889, S.Murbeck; lecto: LD, designated here; iso: S.


Figure 113. Juncus articulatus subsp. articulatus × Juncus bulbosus. A–C, habit and 

inflorescences; D, tepals and stamens; E, perianth and degenerated ovary. Scale bars: 

A–C = 5 cm; D, E = 1 mm. Drawn by Ö.Nilsson. Reproduced with permission, from 

Ö.Nilsson & S.Snogerup, Bot. Not. 125: 204, fig. 75 (1972). © Lund Botanical Society. 

265

266 


Figure 114. Juncus acutiflorus × Juncus articulatus subsp. articulatus. A, habit and 

inflorescences; B, flower head enlarged by continued flowering; C, tepals and stamens; 

D, perianth and capsule. Scale bars: A = 5 cm; B = 3 mm; C, D = 1 mm. Drawn by 

Ö.Nilsson. Reproduced with permission, from Ö.Nilsson & S.Snogerup, Bot. Not. 125: 205, 

fig. 76 (1972). © Lund Botanical Society.


Juncus anceps × lampocarpus B. [unranked] frisicus Buchenau ex Asch. & Graebn., Syn. 

Mitteleur. Fl. 2 (2): 479 (1904), nom. inval. 

Juncus ×frisicus Dörfl., Sched. Herb. Norm. no. 4783 (1907). Authentic material: [Germany] Borkum, 1894, 

F.Wirtgen; n.v.; Baltrum, 1900, F.Buchenau; n.v. 

Juncus ×livetianus H.Lév., Bull. Soc. Bot. France 53: 535 (1906) 

T: [France] Sarthe, Livet, bord du ruisseau de Bécherel, 28 Aug 1906, H.Léveillé; syn: n.v. 


An almost completely seed-sterile hybrid. It is known to occur in several areas of common 

occurrence of the two species (Denmark, Sweden, France, Germany, Bosnia-Herzegovina). 

2n = 60, fide M.Zandee, Proc. Kon. Ned. Akad. Wetensch., C, 84: 250 (1981). 10: DEN, 

SWE. 11: GER. 12: FRA. 13: YUG. 

10. DENMARK: Jutland, Kirkeflod at Lønne, 21 Aug 1966, B.Øllgaard (W). 

Juncus ×degenianus Boros, Magyar Bot. Lapok. 19 (1920): 41 (1922) = Juncus articulatus L. 

× Juncus subnodulosus Schrank 

T: [Hungary, Budapest area]in paludosis rivi "Alagi majori patak" inter pagos Rákospalota et Alag, 23 Jun 

1917, A.Boros; holo: BP, n.v. 

Material not seen; the apparently extremely rare hybrid is reported for the above single site. 

Juncus ×rusguniensis Trab. ex Maire, Fl. Afr. Nord 4: 302 (1957) [nom. valid., a reference to 

Buchenau, in H.G.A.Engler, Pflanzenr. (iv.36) 25: 192 (1906) was given] = Juncus 

articulatus L. × Juncus fontanesii J.Gay ex Laharpe 

T: Algeria, Massauta, Fort de l'Eau, Apr 1913, L.C.Trabut 1417; syn: P. 

Occasionally between parental plants. It is seed-sterile, usually with developed capsules. 

Hybrids between type subspecies of both parents are known from Algeria, Portugal and 

France. Hybrid between Juncus articulatus subsp. articulatus and Juncus fontanesii subsp. 

pyramidatus are not rare in Turkey, e.g., Adana, between Gürümze and Suphandere, 

P.H.Davis 19592 (ANK, E); Içel, Mersin, E.Hennipman & al. 1334 (L). Also seen from 

Greece, Cyprus, Lebanon and Palestine. Hybrids with subsp. kotschyii have been found in Iran, 

E Sanandag, 6000 ft, Furse 3347 (K) and Pakistan, Quetta, Urak, 2000 m, R.R.Stewart 617 (K). 

Juncus anatolicus Snogerup × Juncus articulatus L. 

Intermediate between the parents, completely seed-sterile, propagating vegetatively and 

locally abundant in hard-grazed subalpine meadows. Known from Turkey, Bursa, Ulu Dagh, 

Sarialani, E.Leick 275 (ISTF); Ulu Dagh, N of the summit, 2000 m, S.Snogerup & 

B.Snogerup 4762 (LD). 

Juncus articulatus L. × Juncus atratus Krock. 

Intermediate between the parents, completely seed-sterile. Only known from Turkey [Van, 

Çaldiran, 2200 m, F.Sorger & Buchner 82–60–12 (W)]. 

Juncus articulatus L. × Juncus striatus Schousb. ex E.Mey. 

Similar to Juncus striatus. It exhibits a considerable vegetative growth, is completely seedsterile 

and its capsule is imperfectly developed. Reported to occur in France, Spain, Greece 

and the European part of Turkey [e.g., France, Charente, Villefagnan, E.Contré [Soc. Fr. 

Éch. Pl. Vasc.] 4710 (L, P), as Juncus ×valbrayi H.Lév.; Turkey-in-Europe, Kirklareri, 

A.Baytop (ISTE 32445)]. 

267

268 


Juncus ×fallax Trab., in J.A.Battandier & L.C.Trabut, Fl. Algérie, ed. 2, 86, tab. 2 (1895) 

= Juncus fontanesii J.Gay ex Laharpe × Juncus valvatus Link 

T: Algeria, Constantine, Djebel Ouach, L.C.Trabut; syn: P, n.v. 

Juncus striatus var. macrocephalus Coss. & Durieu, in J.B.G.M.Bory, Expl. Sci. Alger. 2: 269 (1868). 

T: Algeria, Constantine, Mansoura, E.S.-C.Cosson; syn: n.v.; Algiers, Dra-el-Mizan, E.S.-C.Cosson; syn: n.v. 

Rarely between parental taxa, probably sterile but material not seen. Known from North 

Africa. 

Juncus ×donyanae Fern.-Carv., Lagascalia 14(1): 117 (1986) = Juncus acutiflorus Ehrh. ex 

Hoffm. subsp. rugosus (Steud.) Cout. × Juncus striatus Schousb. ex E.Mey. 

T: [Spain] Huelva, El Rocío, Reserva Biológica de Doñana, Caño de la Raya, 18 Jun 1978, E.Valdés-Bermejo 

& S.Castroviejo; holo: FCO10375; iso: FCO22170. 

Stems striate at the lower part, rugose upwards. Cauline leaves 2–3, rugose. Inflorescence 

with 6–23 semiglobose or globose heads, each with (6–) 10–26 flowers. Tepals unequal; 

inner tepals 3–3.5 mm long; outer tepals 2.6–3 mm long. Anthers 1–12 mm long; filaments 

0.35–0.5 mm long. Capsules 3.4–3.7 mm long. Known only from SW Spain. Sterile. 

Juncus fontanesii J.Gay ex Laharpe × Juncus striatus Schousb. ex E.Mey. 

A rare hybrid characterized by high sterility, leaf surface with delicate longitudinal lamellae, 

and an elongated subterranean creeping stem. It is known to occur in Algeria and Greece. 

13: GRE. 20: ALG. 

13. GREECE: Epiros, Preveza, Archangelos, 29 May 1971, E.Stamatiadou 12462 (ATH). 20. ALGERIA: 

Algiers, 1859–1861, A.Lerou (MANCH). 

Juncus ×valbrayi H.Lév., Bull. Soc. Bot. France 54: 518 (1907) = Juncus anceps Laharpe 

× Juncus acutiflorus Ehrh. ex Hoffm. 

Juncus ×sylvaticoides H.Lév., Bull. Géogr. Bot., sér. 4, 25: 48 (1917), nom. illeg. T: [France] Sarthe, Livet, 

Valbray, près du ruisseau de Bécherel, Sep 1907, H.Léveillé; syn: n.v. 

Although the existence of the hybrid is plausible, we have not seen any material referable to 

it; up to now reported only from France. 

Juncus articulatus L. × Juncus holoschoenus R.Br. 

Rarely at contact between parental taxa, sterile. Known from Australia and New Zealand. 

50: NSW. 51: NZN. 

50. NEW SOUTH WALES: Majors Creek, Apr 1968, Flack s.n. (NSW99369). 51. NORTH IS.: Standish 

Road, E of Stratford, Taranaki, [R.]Mason 4219 (NSW, CHR); near Bell Block, Taranaki, [R.]Mason 4262 

(NSW, CHR). 

Juncus articulatus L. × Juncus thompsonianus L.A.S.Johnson 

Found at a single site in New South Wales among parental plants; probably sterile. 50: NSW. 

50. NEW SOUTH WALES: S Tablelands, Junction of Wragges and Pipers Creek, Kosciusko National Park, 

17 May 1976, L.A.S.Johnson 8282 (NSW). 

A hybrid between J. acutiflorus and J. subnodulosus was reported from Isle Wight, 1924, 

E.Drabble (BM), fide E.Drabble & J.W.Long, Rep. Brit. Exch. Club 9: 734–757 (1932); 

C.A.Stace, Hybridization Fl. Brit. Isl. 465 (1975). The voucher specimen is difficult to 

identify but the occurrence of the hybrid is rather improbable.


Doubtful or unclear taxa 

Juncus isthmiacus Neck., Delic. Gallo-Belg. 1: 168 (1768) 

T: not designated, nor selected here. 

Necker referred back to Linnaean phrase-names associated with the binomial Juncus 

articulatus L. However, the reference does not cover the whole mixture of elements covered 

by the binomial, and perhaps one of the syntypes may be given preference because of 

Necker's note "icon bona" after the citation Gramen junceum aquaticum B. Prodr. 12 

[= C.Bauhin, Prodr. 12 (1620), which is var β of Linnaeus' J. articulatus]. The identification 

of the syntype remains to be done; the name J. isthmiacus may require rejection if it is not 

equated with J. articulatus. 

Juncus annuus Krock., Fl. Siles. 1: 566 (1787) 

T: [Poland, Sląsk, near Wrocław] icon, in A.J.Krocker, Fl. Siles. 1: [566–567] tab. 51 (1787); syn. 

The description and the figure point to Juncus bulbosus but some features of both do not 

exclude the Juncus bufonius group. The name is published as Juncus annuus Milleri (not 

'noster' as in other new taxa) but the reference to Miller was not traced to the original source 

(perhaps a reference to a phrase-name including Juncus annuus). 

Juncus nigricans Wolff ex Schreb., in A.F.Schweigger & F.Körte, Fl. Erlang. 149 (1811) 

T: [Germany] Erlangen, J.C.D.Schreber; syn: n.v. 

Juncus schoenoides Mérat, Nouv. Fl. Env. Paris 2(2): 73 (1831), n.v. 

T: [Paris] n.v. [probably a plant infested by Livia juncorum] 

Juncus divaricatus Wolff, in F.X.Heller, Fl. Wirceb. 1: 28 (1810), n.v. 

T: [Germany, Würzburg region]; syn: L904.144–289 

The type specimen belongs to J. subnodulosus but the publication itself was not seen during 

this study, and neither the validity nor type locality statement were checked. 

Juncus acutiflorus var. minor Hook., Fl. Bor.-Amer. 2: 190 (1838), nom. illeg. 

T: Canada, Richardson; syn: BM, K, n.v.; Mrs Sheppard; syn: n.v. 

Juncus polycephalus var. pumilus Hook., Fl. Bor.-Amer. 2: 190 (1838). 

T: Rocky Mountains, J.Drummond; holo: n.v. 

Juncus wulfenii Miel. ex Saut., Flora 32: 666 (1849), nom. inval. [not accepted by its 

author]. 

Authentic material: Austria, Tofern in Grossarl, M.Mielichhofer, n.v. [probably J. alpinoarticulatus] 

Juncus olympicus Schott, Nyman & Kotschy, Analect. Bot. 3 (1854) 

Juncus lampocarpus [unranked] olympicus Asch. & Graebn., Syn. Mitteleur. Fl. 2 (2): 478 (1904). 

T: [Turkey, Ulu Dagh] in Olympo Bithyniae, [collector not given]; holo: n.v. [not found at W] 

According to the description, the name might belong to a depauperate form of J. anatolicus, 

J. articulatus or their hybrid; the first alternative would threaten the name J. anatolicus. 

Juncus nitidus Phil., Fl. Atacam. 52 (1860) 

T: [Chile] prope Cachinal de la Sierra, R.A.Philippi; holo: SGO, n.v. 

269

270 


Juncus lampocarpus a. [unranked] salinus Schur, Enum. Pl. Transsilv. 685 (1866) 


Juncus digeneus Borbás, [Flor. Adat. Roripakra] Értek. Természettud. Köréb. Magyar. Tud. 

Akad. 9: 15 (1879), n.v. 

T: n.v. 

Published as a binomial for plants believed to represent a hybrid between J. conglomeratus 

and J. thomasii, an extremely improbable hybrid combination. Material not seen. Interpreted 

as a dwarf form of J. thomasii by M.Raciborski, Sprawozd. Kom. Fizyjogr. Akad. Umiej. 

22(2): [p. separ.] 17 (1888). 

Juncus supinus b. erectus E.Lehm., Arch. Naturk. Liv.-Ehst.-Kurlands, Ser. 2, Biol. Naturk. 

11: 176 (1895) 

T: n.v. 

Perhaps a variant of J. bulbosus. 

Juncus nebrodensis Lojac., Fl. Sicul. 3: 161 (1909) 

T: [Sicily] Nebrodi, M.Scalone; syn: n.v.; Malpasso, V.Tineo; syn: n.v.; Madonie; syn: n.v.; M.Lojacono 

Pojero, Fl. Sicul. 3: tab. 13, fig. 2 (1909), as J. affinis; syn. 

It might belong near J. alpinoarticulatus but material not seen. 

Juncus nodifolius Larrañaga, in Anonymous, Escritos Dámaso Antonio Larrañaga 2: 138 

(1923) 


The description points to an affinity with sect. Ozophyllum. 

Juncus articulifolius Larrañaga, in Anonymous, Escritos Dámaso Antonio Larrañaga 2: 138 

(1923) 


The description points to an affinity with sect. Ozophyllum.


APPENDIX I 

List of taxa with lectotypes and neotypes designated in the present volume (by J.Kirscher, if 

not otherwise stated in the text). 

Juncus sect. Articulati T.Marsson 151 

Juncus sect. Graminifolii Engelm. 27 

Juncus ×murbeckii Sagorski 264 

Juncus abortivus Chapm. 165 

Juncus acuminatus f. sphaerocephalus F.J.Herm. 217 

Juncus alatus Franch. & Sav. 145 

Juncus alpigenus K.Koch 182 

Juncus apiculatus Adamson 37 

Juncus aristiflorus Clairv. 249 

Juncus asper Engelm. 171 

Juncus atratus Krock. 178 

Juncus biglumis L. 112 

Juncus brachystigma Sam. 95 

Juncus caespiticius E.Mey. 38 

Juncus canadensis var. brachycephalus Engelm. 176 

Juncus canadensis var. brevicaudatus Engelm. 174 

Juncus canadensis var. longicaudatus Engelm. 173 

Juncus capensis f. depauperatus Buchenau 36 

Juncus capensis subsp. geniculatus Buchenau 37 

Juncus capensis var. latifolius E.Mey. 31 

Juncus caudatus Chapm. 173 

Juncus concinnus D.Don 80 

Juncus elegans Royle ex Sam. 80 

Juncus elliottii Chapm. 221 

Juncus fasciculatus Schrank 161 

Juncus fockei Buchenau 185 

Juncus fuscoater subsp. arthrophyllus Brenner 241 

Juncus holoschoenus R.Br. 185 

Juncus kingii Rendle 95 

Juncus lagenarius Gay ex Laharpe 235 

Juncus lampocarpus Ehrh. ex Hoffm. 151 

Juncus lampocarpus var. lindhardii Wiinst. 251 

Juncus leucanthus Royle ex D.Don 92 

Juncus longistamineus A.Camus 133 

Juncus melananthos Rchb. 178 

Juncus maximowiczii Buchenau 134 

Juncus nikkoensis Satake 228 

Juncus obtusiflorus Ehrh. ex Hoffm. 167 

Juncus planifolius R.Br. 41 

Juncus prismatocarpus subvar. pluritubulosus Buchenau 149 

Juncus rariflorus Hartm. 243 

Juncus rariflorus var. obtusatus Hartm. 243 

Juncus scirpoides var. compositus R.M.Harper 208 

271

272 


Juncus scirpoides var. meridionalis Buchenau 201 

Juncus subcuneatus var. latifolius Adamson 33 

Juncus subnodulosus Schrank 167 

Juncus sulcatus Hochst. 36 

Juncus supinus Moench 161 

Juncus viviparus Relhan 161 

Juncus xiphioides var. triandrus Engelm. 140

MAPS 

Number in brackets refers to the page on which the taxon is described. 

273

174. Juncus acutus subsp. acutus (6) 

175. Juncus acutus subsp. leopoldii (8) 

274 

SPECIES PLANTARUM — FLORA OF THE WORLD (2002)

176. Juncus littoralis (8) 177. Juncus heldreichianus subsp. heldreichianus (12) 

178. Juncus heldreichianus subsp. orientalis (12) 179. Juncus socotranus (14) 

275

180. Juncus maritimus (14) 

181. Juncus rigidus (16) 

276 


182. Juncus kraussii subsp. kraussii (20) 183. Juncus kraussii subsp. australiensis (20) 

184. Juncus kraussii subsp. austerus (22) 185. Juncus roemerianus (22) 

186. Juncus cooperi (25) 187. Juncus sparganiifolius (29) 

277

278 


188. Juncus engleri (30) 189. Juncus lomatophyllus (31) 

190. Juncus dregeanus subsp. dregeanus (33) 191. Juncus dregeanus subsp. bachitii (34) 

192. Juncus sonderianus (34) 193. Juncus capensis (36)

194. Juncus caespiticius (38) 195. Juncus meianthus (38) 

196. Juncus antarcticus (40) 

279

197. Juncus planifolius (41) 

280 


198. Juncus regelii (41) 199. Juncus howellii (42)

200. Juncus repens (42) 201. Juncus cyperoides (45) 

202. Juncus filipendulus (46) 

281

203. Juncus marginatus (46) 

282 


204. Juncus falcatus subsp. falcatus (49) 

205. Juncus falcatus subsp. sitchensis (50) 206. Juncus covillei var. covillei (51) 

283

284 


207. Juncus covillei var. obtusatus (51) 208. Juncus prominens (51) 

209. Juncus orthophyllus (53) 210. Juncus longistylis (53) 

211. Juncus macrophyllus (55)

212. Juncus capitatus (59) 

213. Juncus triformis (60) 214. Juncus leiospermus var. leiospermus (63) 

285

286 


215. Juncus leiospermus var. ahartii (63) 216. Juncus kelloggii (64) 

217. Juncus luciensis (64) 218. Juncus tiehmii (66) 

219. Juncus capillaris (66) 220. Juncus bryoides (67)

221. Juncus uncialis (67) 222. Juncus hemiendytus var. hemiendytus (68) 

223. Juncus hemiendytus var. abjectus (70) 224. Juncus scabriusculus (70) 

225. Juncus obliquus (71) 226. Juncus stenopetalus (71) 

287

288 


227. Juncus rupestris (72) 228. Juncus pictus (72) 

229. Juncus cephalotes (73) 230. Juncus concinnus (80) 

231. Juncus khasiensis (81) 232. Juncus gracilicaulis (81)

233. Juncus grisebachii (82) 234. Juncus chrysocarpus (82) 

235. Juncus clarkei (83) 236. Juncus hydrophilus (84) 

237. Juncus crassistylus (84) 238. Juncus fimbristyloides (86) 

289

290 


239. Juncus ochraceus (86) 240. Juncus allioides (89) 

241. Juncus membranaceus (89) 242. Juncus glaucoturgidus (90) 

243. Juncus leucanthus (92) 244. Juncus spectabilis (94)

245. Juncus cephalostigma (94) 246. Juncus brachystigma (95) 

247. Juncus kingii (95) 248. Juncus trichophyllus (96) 

249. Juncus thomsonii (96) 

291

250. Juncus triglumis subsp. triglumis (98) 

251. Juncus triglumis subsp. albescens (99) 

292 


252. Juncus leucomelas (99) 253. Juncus benghalensis (101) 

254. Juncus perpusillus (101) 255. Juncus concolor (102) 

256. Juncus przewalskii (102) 257. Juncus sherei (104) 

293

JUNCACEAE (Juncus) - Species Plantarum Programme

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