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bolboschoenus fluviatile, river bulrush, river tuber-bulrush, scirpe fluviatile

bulrush, tuberous bulrush, tuberous bulrushes

Habit Herbs, perennial, cespitose or not, rhizomatous.
Culms

100–200 cm × 5–15 mm.

solitary or not, cormose at base [except eastern Asian Bolboschoenus planiculmis (F.

Leaves

sheaths reaching to middle of culm or higher, fronts convex (to concave) and papery at mouth, veins reaching apex, very rarely diverging leaving triangular, veinless, membranous area;

widest blade 7–22 mm wide.

basal and cauline;

sheaths tubular;

ligules absent;

blades flat or V-shaped in cross section, prominently keeled on abaxial surface, distally trigonous.

Inflorescences

subumbellate, with all or most spikelets solitary or in clusters of 2–3(–8) on 4–12 rays, rays not exceeding 10 cm;

involucral bracts that surpass inflorescences 3–6, widest bract 4–15 mm wide.

terminal, subumbellate, subcorymbose, or capitate;

spikelets to 80, mostly 4–10 mm diam.;

involucral bracts surpassing inflorescence 1–5, spreading or proximal erect, leaflike.

Spikelets

10–40, ovoid to lanceoloid, 10–25 × 6–10 mm, base cuneate to rounded;

scales often loosely imbricate, orange-brown to stramineous, usually obscurely lineolate-spotted, 7–10 × 3–4 mm, membranous and translucent, apex 2-fid 0.5–1 mm deep, awns fairly stout, 2–3 × 0.5 mm wide at base.

scales 25 or more, spirally arranged, each subtending flower, abaxially puberulent, often glabrescent, tip notched and awned.

Flowers

perianth bristles tightly attached to shed achene, pale brown, very stout, equaling achene;

anthers yellow, 4 mm;

styles 3-fid.

bisexual;

perianth of 3–6 bristles, straight or curved, shorter to slightly longer than achene, retrorsely spinulose;

stamens 3;

styles linear, 2–3-fid, base persistent.

Achenes

grayish or dark brown, often in patches, obovoid to obpyriform, all nearly equilaterally markedly trigonous to slightly compressed, angles equally rounded, 3.8–5.5 × 2–2.9 mm, apex rounded, beak 0.2–0.8 mm, surface rather dull, exocarp cells usually not evident at 20X; in cross section exocarp much thinner than mesocarp and its cells very small, isodiametric;

achene specific gravity greater than water.

biconvex to trigonous, 2.3–5.5 mm, smooth.

Schmidt

) T.

v

.

Egerova

], sharply trigonous.

2n

= 94.

Bolboschoenus fluviatilis

Bolboschoenus

Phenology Fruiting summer.
Habitat Fresh shores, inland marshes, coastal estuaries
Elevation 0–2100 m (0–6900 ft)
Distribution
from FNA
AL; AZ; CA; CO; CT; DE; IA; ID; IL; IN; KS; MA; MD; ME; MI; MN; MO; MT; ND; NE; NH; NJ; NY; OH; OR; PA; SD; TN; UT; VA; VT; WA; WI; AB; BC; MB; NB; ON; QC; SK; Asia (Japan); Australia; Pacific Islands (New Zealand)
[WildflowerSearch map]
from USDA
Nearly worldwide
Discussion

Bolboschoenus fluviatilis frequently forms dense, monospecific, often entirely vegetative stands, and it is more common than recorded because vegetative colonies are often overlooked (E. W. Chester and B. E. Wofford 1992). The only record for Alabama is an 1870 collection from the East Fowl River in the Mobile Delta, where the species has not been collected since. It was intentionally introduced into New Hampshire (D. J. Padgett and G. E. Crow 1993). The report from New Mexico by M. L. Fernald (1950) cannot be confirmed because no specimen is known.

Putative hybrids with Bolboschoenus maritimus occur in California. Bolboschoenus novae-angliae probably originated from B. fluviatilis × B. robustus (J. Browning et al. 1995). Introgression from B. maritimus and/or B. robustus is suggested by the larger exocarp cells (evident in surface view) in some North American plants. The Eurasian B. yagara (Ohwi) Y. C. Yang & M. Zhan differs from B. fluviatilis in its narrower leaves and smaller achenes.

(Discussion copyrighted by Flora of North America; reprinted with permission.)

Species 6–15 (5 in the flora).

Bolboschoenus comprises at least six to as many as fifteen species, depending on the treatments of the B. maritimus and B. fluviatilis complexes and taxa of probable hybrid origin (J. Browning et al. 1995). Treatments of the genus published prior to J. Browning and K. D. Gordon-Gray (1993) and J. Browning et al. (1995) are unsatisfactory because of reliance on highly variable, nondiagnostic characters. The pericarp anatomy of the achene and the persistence of perianth bristles on shed achenes provide especially critical but seldom-used taxonomic characters. Detailed revisions utilizing these characters have been undertaken only in a few areas of the world, leaving many problems that cannot be resolved until a worldwide revision is completed. Pericarp anatomy is easily observed by examining hand-sections with a dissecting microscope. Achenes of the halophytic species B. maritimus and B. robustus have caducous bristles, and exocarp (epidermal) cells are lighter than water, greatly enlarged, and air-filled so that the achenes float for many days. In contrast, in the freshwater species B. fluviatilis and B. glaucus the bristles remain attached to the achenes and exocarp cells are very small; the achenes are denser than water and quickly sink (J. Browning et al. 1995).

Bolboschoenus is further usually characterized by culms that are often distally scabrous or papillose on margins, and leaf blades that are antrorsely spinulose or papillose on margins and often abaxially on midribs. Those characteristics, in addition to the presence of durable corms at culm bases, noded culms (i.e., some leaves cauline), the complete absence of ligules, the abaxially keeled leaf blades, and puberulent spikelet scales that are apically 2-fid and prominently awned are diagnostic of all species worldwide except the eastern Asian B. planiculmis (F. Schmidt) T. V. Egorova.

In recent years upon the dismemberment of Scirpus in the broad sense, the perponderance of evidence supports the recognition of Bolboschoenus as distinct from Schoenoplectus (C. D. Adams 1994c; J. Browning and K. D. Gordon-Gray 1993; J. Browning et al. 1995; J. J. Bruhl 1995; P. Goetghebeur and D. A. Simpson 1991; E. Hayasaka and H. Ohashi 2002; Z. Hroudová et al. 1998; T. Koyama 1980; A. A. Oteng-Yeboah 1974b). Bolboschoenus was included within Schoenoplectus by K. A. Lye (1971) on the basis of similarities in embryo morphology and by M. T. Strong (1993) on the basis of similarities in embryo, leaf, and inflorescence morphology, as well as the intermediate structure of the eastern Asian B. planiculmis. The similarities between Bolboschoenus and Schoenoplectus in leaves and inflorescences as pointed out by M. T. Strong (1993) appear to be superficial; and they do not include other features usually characteristic of Bolboschoenus as given above. Although B. planiculmis is similar to some species of Schoenoplectus in having all leaves basal, leaf blades trigonous, involucral bracts solitary and trigonous, and in lacking hard corms at the shoot bases, it clearly belongs to Bolboschoenus on the basis of the absence of ligules, as well as similarities in the achenes and floral scales. The vegetative morphology of B. planiculmis is probably an adaptaton to its highly aquatic, tidal habitat (T. Koyama 1980; E. Hayasaka and H. Ohashi 2002).

In North America the following putative hybrids are known from regions of sympatry of the parental species: 1) Bolboschoenus maritimus × B. robustus, common along both Atlantic and Pacific coasts; 2) B. maritimus × B. glaucus, local in California and Idaho; 3) B. fluviatilis × B. robustus [= ×B. novae-angliae], uncommon on the Atlantic Coast; 4) B. fluviatilis × B. maritimus, uncommon and local in California. Putative hybrids are often sterile; achenes that are formed are often empty or contain abnormal embryos.

Species of Bolboschoenus are often ecologically dominant in wetlands and are valuable food and habitat for wildlife (H. A. Kantrud 1996). Some species are cultivated in wildlife refuges in the United States and are important rice-field weeds in California.

In this treatment, spikelet and floral scale measurements are for nonflattened structures and exclude awns; awn widths are often difficult to measure: floral scale apices frequently are so worn that the lobes are not evident and thus apices appear to taper into awns, and scales often split distally making it difficult to determine the position of the base of the awn; floral scale translucency is best determined with a dissecting microscope by placing a shiny needle beneath the scale; achene lengths include the beak and, if present, the persistent remnant of the style base; culm widths given herein are measured on the middle third of the culm; and inflorescence descriptions are for expanded inflorescenses.

(Discussion copyrighted by Flora of North America; reprinted with permission.)

Key
1. Inflorescences with all spikelets sessile or not more than 1/2 on evident branches; perianth bristles not persistent on shed achene or 1–few bristles weakly persistent; achenes biconvex or compressed-trigonous with low, rounded abaxial angle; achene epidermal cells appearing greatly enlarged in cross section, 2–3+ times deeper than wide; styles 2–3-fid; habitats brackish to saline.
→ 2
1. Inflorescences with all or most spikelets on evident, elongated branches; perianth bristles tightly attached to shed achene, rarely some or all falling separately; achenes equilaterally trigonous to compressed-trigonous or rarely some biconvex; achene epidermal cells appearing small in cross section, isodiametric or very rarely enlarged and to 3 times deeper than wide (in B. novae-angliae); styles mostly 3-fid; habitats fresh to slightly brackish.
→ 3
2. Floral scales hyaline, membranous, awns 0.25 mm wide at base; anthers yellow, rarely orange; styles 2-fid, rarely 3-fid; achenes usually biconvex, rarely compressed trigonous; leaf sheath fronts usually membranous at mouth, veins diverging proximal to summit leaving triangular, veinless area.
B. maritimus
2. Floral scales nearly opaque, usually papery, awns 0.5 mm wide at base; anthers brownish orange; styles 3-fid or a few 2-fid; achenes mostly trigonous; leaf sheath fronts usually papery at mouth, veins reaching nearly to summit.
B. robustus
3. Widest leaf blade 2–6 mm wide; leaf sheath fronts membranous at mouth, veins diverging proximal to summit leaving acutely triangular veinless area; spikelets 3–5 mm diam.; scale awns slender, 0.25 mm wide at base; achenes 2.5–3.3 mm, beak minute.
B. glaucus
3. Widest leaf blade 7–22 mm wide; leaf sheath fronts mostly papery at mouth, veins nearly reaching summit; spikelets 5–10 mm diam.; scale awns stout, to 0.5 mm wide at base; achenes 3–5.5 mm, beak distinct, 0.1–0.8 mm.
→ 4
4. Achenes all nearly equilaterally to slightly compressed, markedly trigonous, 3.8–5.5 mm; perianth bristles all firmly attached to shed achenes; fresh water habitats, mostly inland.
B. fluviatilis
4. Achenes variably compressed, markedly to obscurely trigonous or some biconvex in same spikelet, 3–4.3 mm; perianth bristles weakly attached to shed achenes or many caducous; slightly brackish habitats, coastal estuaries.
B. novae-angliae
Source FNA vol. 23, p. 42. FNA vol. 23, p. 37. Author: S. Galen Smith.
Parent taxa Cyperaceae > Bolboschoenus Cyperaceae
Sibling taxa
B. glaucus, B. maritimus, B. novae-angliae, B. robustus
Subordinate taxa
B. fluviatilis, B. glaucus, B. maritimus, B. novae-angliae, B. robustus
Synonyms Scirpus maritimus var. fluviatilis, Scirpus fluviatilis, Schoenoplectus fluviatilis Scirpus section B.
Name authority (Torrey) Soják: Cas. Nár. Mus., Odd. Prír. 141: 62. (1972) (Ascherson) Palla: in W. D. J. Koch et al., Syn. Deut. Schweiz. Fl. ed. 3, 3: 2531. (1905)
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