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amphibious bistort, r enouée amphibie, water knotweed, water smartweed

knotweed

Habit Plants perennial, 2–12 dm in terrestrial plants, to 30 dm in some aquatic plants; roots also sometimes arising from proximal nodes; rhizomes or stolons usually present. Trees, shrubs, vines, or herbs, perennial or annual, homophyllous (heretophyllous in some species of Polygonum); root fibrous or a solid taproot, rarely tuberous.
Stems

prostrate to ascending or erect, simple or branched, ribbed, glabrous or strigose to hirsute.

usually prostrate to erect, sometimes scandent, not scapose, rarely with recurved spines (some species of Persicaria), glabrous or pubescent, sometimes glandular;

nodes usually swollen;

branches free (adnate to stems distal to nodes and appearing to arise internodally in Polygonella);

tendrils absent (except in Antigonon and Brunnichia).

Leaves

ocrea tan to dark brown, cylindric or flared distally, 5–50 mm, chartaceous or, sometimes, foliaceous distally, base inflated, margins truncate to oblique, glabrous or ciliate with hairs 0.5–4.5 mm, surface glabrous or appressed-pubescent to hirsute, not glandular-punctate;

petiole 0.1–3(–7) cm, glabrous or appressed-pubescent to hirsute, leaves sometimes sessile;

blade without dark triangular or lunate blotch adaxially, ovate-lanceolate to elliptic or oblong-lanceolate, 2–15(–23) × 1–6(–8) cm, base usually tapered to acute or rounded, rarely cordate, margins antrorsely scabrous, apex acute to acuminate, faces glabrous or sparingly strigose, midveins glabrous or strigose, not glandular-punctate.

deciduous (persistent in Coccoloba and sometimes more than 1 year in Antigonon and Polygonella), basal or basal and cauline, rarely cauline only, mostly alternate;

ocrea present, persistent or deciduous, cylindric to funnelform, chartaceous, membranous, coriaceous, or, rarely, foliaceous or partly so;

petiole present or absent, rarely articulate basally (Fagopyrum, Fallopia, Polygonella, Polygonum), rarely with extrafloral nectaries (Fallopia, Muehlenbeckia);

blade simple with entire margins, rarely undulate or lobed.

Inflorescences

terminal, ascending to erect, uninterrupted or interrupted proximally, 10–150 × 8–20 mm;

peduncle 10–50 mm, glabrous or strigose to hirsute, often stipitate-glandular;

ocreolae overlapping except sometimes proximal ones, margins ciliate with bristles to 1 mm.

terminal or terminal and axillary, spikelike, racemelike, paniclelike, cymelike, or, rarely, capitate, comprising simple or branched clusters of compound inflorescences;

bracts absent;

peduncle spreading to erect, sometimes absent;

clusters of flowers subtended by connate bracteoles forming persistent membranous tube (ocreola), awnless.

Pedicels

ascending, 0.5–1.5 mm.

Flowers

bisexual or functionally unisexual, some plants having only staminate flowers, others with only pistillate flowers, 1–3(–4) per ocreate fascicle, heterostylous;

perianth roseate to red, glabrous, not glandular-punctate, slightly accrescent;

tepals 5, connate ca. 1/3 their lengths, obovate to elliptic, 4–6 mm, veins prominent, not anchor-shaped, margins entire, apex rounded to acute;

stamens 5, included or exserted;

anthers pink or red, elliptic;

styles 2, included or exserted, connate 1/2–2/3 their length.

usually bisexual, sometimes bisexual and unisexual on same plant, rarely unisexual only, 1–20+ per ocreate fascicle, often with stipelike base distal to articulation;

perianth often accrescent in fruit, often greenish, white, pink, yellow, red, or purple, usually unwinged and unkeeled (winged or, sometimes, keeled in Fallopia, rarely keeled in Polygonum), campanulate or urceolate, sometimes membranous, indurate, or fleshy in fruit, rarely developing raised tubercles proximally (Rumex), glabrous or pubescent, sometimes glandular or glandular-punctate;

tepals 2–6, usually in 2 whorls, distinct or connate proximally and forming tube, petaloid or sepaloid, monomorphic or dimorphic;

nectary a disk at base of ovary or glands associated with bases of filaments;

stamens usually (1–)6–9, staminodes rarely present;

filaments distinct, or connate basally and sometimes forming staminal tube, free or adnate to perianth tube;

pistils (2–)3(–4)-carpellate;

ovary 1-locular (sometimes with vestigial partitions proximally);

ovule 1, orthotropous or, rarely, anatropous, placentation basal or free-central;

styles 1–3, erect to spreading or recurved, distinct or connate proximally;

stigmas peltate, capitate, fimbriate, or penicillate.

Achenes

included, dark brown, biconvex, (2–)2.2–3 × (1.5–)1.8–2.6 mm, shiny or dull, smooth or minutely granular.

yellowish, brown, red, or black, homocarpic (sometimes heterocarpic in Polygonum), winged or unwinged, usually 2–3-gonous, sometimes discoid, biconvex, or spheroidal, rarely 4-gonous.

Seeds

embryo usually straight or curved, rarely folded.

2n

= 66, 132.

Persicaria amphibia

Polygonaceae subfam. polygonoideae

Phenology Flowering Jun–Sep.
Habitat Shallow water, shorelines of ponds and lakes, banks of rivers and streams, moist prairies and meadows
Elevation 0-3000 m (0-9800 ft)
Distribution
from FNA
AK; AR; AZ; CA; CO; CT; DC; DE; IA; ID; IL; IN; KS; KY; LA; MA; MD; ME; MI; MN; MO; MS; MT; NC; ND; NE; NH; NJ; NM; NV; NY; OH; OK; OR; PA; RI; SC; SD; TN; TX; UT; VA; VT; WA; WI; WV; WY; AB; BC; MB; NB; NL; NS; NT; PE; QC; SK; YT; SPM; Mexico; South America; Ont ; Eurasia; Africa
[WildflowerSearch map]
[BONAP county map]
Mainly temperate regions of North America
Discussion

Persicaria amphibia is widespread in the Northern Hemisphere and naturalized in Mexico, South America, and southern Africa. It is highly polymorphic and the most hydrophytic of the native North American smartweeds (R. S. Mitchell 1976). In recent decades, botanists have tended to follow Mitchell (1968) in recognizing two endemic, intergrading North American varieties. Studies by G. Turesson (1961) and Mitchell (1968, 1976) have shown that phenotypic extremes in the species are part of a cline of nearly continuous morphological variation that is strongly correlated with submergence, but also with some genetic integrity. Formal recognition of varieties is even less tenable when Eurasian elements also are considered.

Aquatic-adapted plants, which bloom in water or are sometimes stranded on land, have been called var. stipulacea (although that epithet may not be the oldest one available for the taxon). They produce ovoid-conic to short-cylindric inflorescences 10–40(–60) mm, prostrate aerial stems, and leaf blades that are glabrous with acute to rounded apices. Terrestrial forms of this ecotype usually are spreading-pubescent and often bear ocreae that are foliaceous, green, and flared distally, characters found only in North American plants (R. S. Mitchell 1968).

Terrestrial-adapted plants, referred to var. emersa, bloom on moist soil and produce short- to elongate-cylindric inflorescences 40–110(–150) mm, spreading or erect aerial stems, and leaf blades that are appressed-pubescent with acute to acuminate apices. They produce ocreae that are entirely chartaceous and not flared distally. Emergent and terrestrial plants of this ecotype exhibit less phenotypic plasticity and a lower frequency of heterostyly than do plants of the aquatic ecotype (R. S. Mitchell 1968).

R. S. Mitchell and J. K. Dean (1978) and H. R. Hinds (2000) recognized var. amphibia, the Eurasian element, as introduced in New York and New Brunswick, respectively. These plants are morphologically intermediate between the North American ecotypes and often indistinguishable from North American plants (Mitchell and Dean).

The Meskwaki and Ojibwa used leaves, stems, and roots as a drug to treat a variety of maladies, the Potawatomi used roots to treat unspecified ailments, and the Lakota and Sioux used plants for food (D. E. Moerman 1998).

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

Genera 28, species ca. 850 (16 genera, 160 species in the flora).

Morphological (K. Haraldson 1978; L.-P. Ronse Decraene and J. R. Akeroyd 1988; Ronse Decraene et al. 2000; Hong S. P. et al. 1998) and molecular (A. S. Lamb Frye and K. A. Kron 2003) data provide support for separation of Persicaria from Polygonum. Further studies are needed to elucidate the relationships of allied genera, particularly Aconogonon, Bistorta, and Koenigia with Persicaria, and Fallopia and Polygonella with Polygonum.

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

Key
1. Tendrils present; plants vines
→ 2
1. Tendrils absent; plants trees, shrubs, or herbs, rarely vinelike shrubs
→ 3
2. Perianths pink to purple or, rarely, white or yellow, membranous; pedicels not 3-winged
Antigonon
2. Perianths green to greenish yellow, indurate; pedicels 3-winged, 1 wing more prominent and becoming greatly expanded in fruit
Brunnichia
3. Plants trees or shrubs; tubes of pistillate flowers becoming fleshy in fruit
Coccoloba
3. Plants herbs, subshrubs, or, rarely, vinelike shrubs; tubes of pistillate flowers rarely becoming fleshy in fruit
→ 4
4. Tepals 6
→ 5
4. Tepals 3, 4, or 5
→ 7
5. Flowers unisexual; outer 3 tepals of pistillate flowers each with apex ending in stout spine
Emex
5. Flowers bisexual or, rarely, unisexual; outer 3 tepals each without apex ending in stout spine
→ 6
6. Achenes winged; inner tepals of fruiting perianths nonaccrescent; stamens (6-)9
Rheum
6. Achenes unwinged; inner tepals of fruiting perianths usually accrescent; stamens 6
Rumex
7. Herbs annual; tepals 3 [4]; stamens (1-)3[-5]
Koenigia
7. Herbs perennial or annual, or shrubs; tepals 4-5; stamens 3-8
→ 8
8. Tepals 4; achenes lenticular, winged; leaves mostly basal
Oxyria
8. Tepals 4 or 5; achenes 3-gonous, discoid, biconvex, spheroidal, or 4-gonous, unwinged or essentially so; leaves cauline or basal and cauline, rarely mostly basal
→ 9
9. Branches adnate to stems, appearing to arise internodally
Polygonella
9. Branches not adnate to stems, not appearing to arise internodally
→ 10
10. Plants shrubs, vinelike; flowers unisexual, tubes of pistillate flowers becoming fleshy in fruit
Muehlenbeckia
10. Plants herbs or, if shrubs, not vinelike; flowers bisexual or, rarely, unisexual, if unisexual then tubes of pistillate flowers not becoming fleshy
→ 11
11. Outer tepals winged or keeled
→ 12
11. Outer tepals unwinged and unkeeled
→ 13
12. Outer tepals winged (keeled in F. ciliondis and, usually, F. convolvulus); ocreae chartaceous, tan to brownish, glabrous or scabrous to variously pubescent, never 2-lobed distally
Fallopia
12. Outer tepals keeled; ocreae often hyaline, silvery, glabrous, 2-lobed distally
Polygonum
13. Leaves mostly basal, some cauline; inflorescences terminal, spikelike; stems simple
Bistorta
13. Leaves cauline; inflorescences terminal and axillary or axillary; stems usually branched, rarely simple
→ 14
14. Achenes strongly exserted; perianths nonaccrescent; tepals distinct
Fagopyrum
14. Achenes included or exserted; perianths accrescent or nonaccrescent; tepals connate to 2/3 their lengths. [15. Shifted to left margin.—Ed.]
→ 15
15. Ocreae often hyaline, silvery, glabrous, 2-lobed distally, often disintegrating into fibers or completely
Polygonum
15. Ocreae chartaceous, usually tan, brown, or reddish, rarely silvery, glabrous or scabrous to variously pubescent, never 2-lobed distally, often tearing with age
→ 16
16. Inflorescences spikelike, paniclelike, or capitate; tepals 4 or 15, connate 1/ 2/ 3 their length (less than 5 their length in P. wallichii); stamens 5-8
Persicaria
16. Inflorescences racemelike or paniclelike; tepals 5, connate ca. 1/ 4 their length; stamens 8
Aconogonon
Source FNA vol. 5, p. 583. FNA vol. 5, p. 479. Author: Craig C. Freeman.
Parent taxa Polygonaceae > subfam. Polygonoideae > Persicaria > sect. Persicaria Polygonaceae
Sibling taxa
P. arifolia, P. bicornis, P. bungeana, P. capitata, P. careyi, P. chinensis, P. glabra, P. hirsuta, P. hydropiper, P. hydropiperoides, P. lapathifolia, P. longiseta, P. maculosa, P. meisneriana, P. minor, P. nepalensis, P. orientalis, P. pensylvanica, P. perfoliata, P. punctata, P. robustior, P. sagittata, P. setacea, P. virginiana, P. wallichii
Subordinate taxa
Aconogonon, Antigonon, Bistorta, Brunnichia, Coccoloba, Emex, Fagopyrum, Fallopia, Koenigia, Muehlenbeckia, Oxyria, Persicaria, Polygonella, Polygonum, Rheum, Rumex
Synonyms Polygonum amphibium, P. amphibia var. emersa, P. amphibia var. stipulacea, P. coccinea, P. hartwrightii, P. muhlenbergia, Polygonum amphibium var. emersum, Polygonum amphibium subsp. laevimarginatum, Polygonum amphibium var. natans, Polygonum amphibium var. stipulaceum, Polygonum coccineum, Polygonum coccineum var. pratincola, Polygonum coccineum var. rigidulum, Polygonum emersum, Polygonum hartwrightii, Polygonum natans
Name authority (Linnaeus) Gray: Nat. Arr. Brit. Pl. 2: 268. (1822) Eaton: Bot. Dict. ed. 4, 30. (1836)
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