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angle stem buckwheat, angle-stem wild buckwheat

buckwheat family

Habit Herbs, erect to spreading, annual, 1–5(–10) dm, tomen-tose to floccose or glabrous, usually grayish. Trees, shrubs, vines, or herbs, perennial, biennial, or annual, homophyllous (heterophyllous in some species of Polygonum), polycarpic (rarely monocarpic in Eriogonum); roots fibrous or a solid or, rarely, chambered taproot, rarely tuberous.
Stems

caudex absent;

aerial flowering stems erect, striated, angled, solid, not fistulose, 0.5–1 dm, tomentose to floccose.

prostrate to erect, sometimes scandent or scapose, solid or hollow, rarely with recurved spines (some species of Persicaria), glabrous or pubescent, sometimes glandular;

nodes swollen or not;

tendrils absent (except in Antigonon and Brunnichia);

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

caudex stems (subfam.

Leaves

basal and cauline;

basal: petiole 0.5–3 cm, mostly floccose, blade oblanceolate to oblong-lanceolate, 1–4(–4.5) × (0.2–)0.5–1(–1.3) cm, tomentose abaxially, floccose or glabrate and grayish or greenish adaxially, margins crenulate;

cauline sessile, blade lanceolate to oblong, 0.5–2 × 0.3–0.8 cm, similar to basal blade.

deciduous (persistent in Coccoloba and sometimes more than 1 year in Antigonon, Eriogonum, Chorizanthe, and Polygonella), basal or basal and cauline, rosulate, mostly alternate, infrequently opposite or whorled;

stipule (ocrea) absent (subfam.

Inflorescences

cymose, open, 5–80 × 10–60 cm;

branches striated, angled, sparsely tomentose to glabrate;

bracts 3, scalelike, 1–3 × 1–3 mm.

terminal or terminal and axillary, cymose and dichotomously or trichotomously branched, or racemose, umbellate, or capitate (subfam.

Peduncles

erect, straight, slender, 1–2 cm, sparsely tomentose to glabrous.

Involucres

turbinate-campanulate to campanulate, 1.5–2.5(–3) × 1.5–2.5(–3), sparsely puberulent;

teeth 5, erect, 0.3–0.6 mm.

Pedicels

present or absent, rarely accrescent (Brunnichia), articulate to flowers.

Flowers

1.5–1.8 mm;

perianth white to rose, without a conspicuous rose-purple spot on each outer tepal, minutely glandular-puberulent;

tepals dimorphic, those of outer whorl elliptic to obovate, sometimes inflated proximally, those of inner whorl narrowly spatulate;

stamens exserted, 2–3 mm;

filaments pilose proximally.

usually bisexual, sometimes bisexual and unisexual on same or different plants, rarely unisexual only, 1–many, often with stipelike base distal to articulation;

perianth persistent, often accrescent in fruit, often greenish, white, pink, yellow, red, or purple, rarely winged or keeled (Fallopia and some species of Polygonum), campanulate to urceolate, sometimes membranous, indurate (Brunnichia and Emex), or fleshy (Coccoloba, Muehlenbeckia, and some species of Persicaria) in fruit, rarely developing raised tubercles proximally (Rumex), glabrous or pubescent, sometimes glandular or glandular-punctate;

tepals 2–6, distinct or connate proximally and forming tube, usually in 2 whorls, petaloid or sepaloid, dimorphic or monomorphic, rarely coriaceous (Lastarriaea), entire, emarginate, or lobed to laciniate apically, rarely awn-tipped (Lastarriaea);

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

stamens (1–)6–9, staminode rarely present;

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

anthers dehiscing by longitudinal slits;

pistil 1, (rudimentary pistil sometimes present in staminate flowers of monoecious or dioecious taxa), (2–)3(–4)-carpellate, homostylous (heterostylous in some species of Fagopyrum and Persicaria);

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 1 per style, peltate, capitate, fimbriate, or penicillate.

Fruits

achenes, included or exserted, yellowish, brown, red, or black, homocarpic (sometimes heterocarpic in Polygonum), winged or unwinged, 2-gonous, 3-gonous, discoid, biconvex, lenticular, rarely 4-gonous or spheroidal, glabrous or pubescent.

Achenes

light brown to brown, 3-gonous, 1–1.5 mm, glabrous.

Seeds

1;

endosperm usually abundant, mealy, development nuclear;

embryo straight or curved, rarely folded.

Eriogonoideae

) tightly compact to spreading and at or just below the soil surface or spreading to erect and above the soil surface, woody;

aerial flowering stems prostrate or decumbent to erect, arising at nodes of caudex branches, at distal nodes of aerial branches, or directly from the root, slender to stout and solid or slightly to distinctly fistulose, rarely disarticulating in ringlike segments (Eriogonum).;

eriogonoideae , possibly vestigial in some perennial species of Chorizanthe) or present (subfam.; eriogonoideae );

or spikelike, racemelike, paniclelike, cymelike, or, rarely, capitate (subfam.; eriogonoideae , rarely absent in Eriogonum), glabrous or pubescent;

peduncle present or absent;

clusters of flowers subtended by involucral bracts or enclosed in typically nonmembranous tubular involucres (subfam.; eriogonoideae ) or subtended by connate bracteoles forming a persistent membranous tube (ocreola) (subfam.

Polygonoideae

), persistent or deciduous, cylindric to funnelform, sometimes 2-lobed (Polygonum), chartaceous, membranous, coriaceous or partially to entirely foliaceous;

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

blade simple, margins entire, occasionally crenulate, crisped, undulate, or lobed, rarely awn-tipped (Goodmania).;

polygonoideae ), comprising simple or branched clusters of compound inflorescences;

bracts absent (subfam.; polygonoideae ), or 2–10+, usually connate proximally or to 1/2 their length, rarely perfoliate, foliaceous or scalelike, margins entire, sometimes awn-tipped (subfam.; polygonoideae ).

Eriogonum angulosum

Polygonaceae

Phenology Flowering year-round.
Habitat Clayey flats and slopes, mixed grassland, saltbush, and chaparral communities, oak and conifer woodlands
Elevation 0-800 m (0-2600 ft)
Distribution
from FNA
CA
[WildflowerSearch map]
[BONAP county map]
Widespread; well represented in the north-temperate zone
[BONAP county map]
Discussion

The name Eriogonum angulosum has been applied to all of the members of its species complex except E. gossypinum. Since the 1950s, the name consistently has been applied to plants with long, exserted stamens and strongly angled stems of the Inner Coast Ranges (Alameda and Contra Costa counties south), the western foothills of the southern Sierra Nevada (Tulare County south), and the Central Valley (San Joaquin County south). The southern edge of the range is the northern foothills of the Transverse Ranges (Ventura and Los Angeles counties). The species can be common and occasionally abundant but rarely is weedy. A mixed collection (with E. gracillimum) from Barstow, San Bernardino County (K. Brandegee s.n., May 1913, UC), and two sheets of the species from San Diego gathered by Susan Stokes apparently in 1895 (B, SD) are discounted as to location.

In late fruit, the bractlets at the base of the pedicel inside the involucres of Eriogonum angulosum often elongate and broaden into oblanceolate segments that fill the involucre. As a result, the involucre appears to have several rows of teeth. This feature may be seen also in E. viridescens, but typically the involucres there appear to have only two or three rows of teeth. This feature is seen rarely in E. maculatum.

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

Genera 48, species ca. 1200 (35 genera, 442 species in the flora).

Monophyly of Polygonaceae is well supported by molecular studies (M. W. Chase et al. 1993; P. Cuénoud et al. 2002; A. S. Lamb Frye and K. A. Kron 2003), which place it sister to Plumbaginaceae. Wood anatomy also supports this relationship (S. Carlquist 2003). Polygonaceae has been divided variously into subfamilies (S. A. Graham and C. E. Wood Jr. 1965; K. Haraldson 1978; J. Brandbyge 1993); two often are recognized based on morphological evidence: Eriogonoideae and Polygonoideae. Recent studies using the chloroplast gene rbcL suggest Eriogonoideae are monophyletic and Polygonoideae are paraphyletic (Lamb Frye and Kron). Generic limits have been much debated, particularly the circumscription of Polygonum in the broad sense. Morphological, cytological, palynological, and anatomical data, although useful in resolving some questions about the circumscriptions and relationships of genera, have not provided a consensus regarding relationships within the family. Pollen types (J. W. Nowicke and J. J. Skvarla 1977) and chromosome numbers (Brandbyge) are diverse. Multiple base chromosome numbers have been documented in some genera, and polyploidy is common. The classification used here generally follows J. L. Reveal (1978) for Eriogonoideae and Haraldson for Polygonoideae.

A characteristic feature of the family is the ocrea, a nodal sheath variously interpreted as an outgrowth of the sheathing base of the petiole, as connate stipules, or as an expanded axillary stipule (S. A. Graham and C. E. Wood Jr. 1965). It is absent in subfamily Eriogonoideae except in some perennial, South American members of Chorizanthe, where it is rudimentary (J. L. Reveal 1978).

The flowers of Polygonaceae have been studied extensively. Most researchers concluded that six tepals is the primitive condition in the family (R. A. Laubengayer 1937), but A. S. Lamb Frye and K. A. Kron (2003) concluded that five tepals is the primitive condition, and that taxa with six or four tepals evolved multiple times within the family. The same molecular study also suggested parallel evolution of growth forms and woodiness. Floral nectaries are useful generic characters in the Polygonoideae (L.-P. Ronse Decraene and J. R. Akeroyd 1988) but their utility in floristic treatments is limited because of their small size. In some genera of Polygonaceae, the outer tepals are connate and form a slender, stipelike hypanthium base above the articulation with the true pedicel. In the Polygonoideae treatments, pedicel refers to the true pedicel plus the stipelike hypanthium above the articulation; in Eriogonoideae, stipe refers only to the stipelike hypanthium base, and pedicel is applied only to the true pedicel.

Members of the family are of relatively minor economic importance (A. N. Steward 1930; S. A. Graham and C. E. Wood Jr. 1965). Fagopyrum has been cultivated for millennia, Coccoloba produce edible fruits, and the petioles of Rheum and leaves of some species of Rumex are edible. Many species have been used as food or medicine or for ceremonies by various tribes of Native Americans. Outside the flora area, the family includes some tropical trees harvested for timber (Coccoloba and Triplaris). Some species of Fallopia, Persicaria, Polygonum, and Rumex are cosmopolitan weeds, and Emex is a pernicious weed in parts of the world. Of the 35 genera in the flora, four are entirely non-native: Emex, Fagopyrum, Muehlenbeckia, and Rheum. Antigonon and Coccoloba are popular ornamentals in the southern part of the flora region. Many species of Eriogonum are planted in rock gardens. Persicaria species are important wetland plants because waterfowl consume their fruits.

Mechanisms for achene dispersal vary greatly. Hooked or awned involucres occur in many genera of subfam. Eriogonoideae. The outer tepals of the perianth expand into wings (Fallopia) or spines (Emex), or the achene has wings (Eriogonum alatum, Oxyria, Rheum, and Rumex), which aid in dispersal by wind or water. The achenes are forcibly tossed from the plant in Persicaria virginiana, and persistent, hooked styles aid in their transport by animals. In Coccoloba, the fleshy tube of the tepals aids in dispersal by birds.

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

Key
1. Ocreae absent; nodes not swollen; flowers usually enclosed in involucres or subtended byinvolucral bracts, these rarely absent (in Gilmania)
subfam. Eriogonoideae
1. Ocreae present, persistent or deciduous; nodes usually swollen; flowers not enclosed in involucres or associated with involucral bracts
subfam. Polygonoideae
Source FNA vol. 5, p. 411. FNA vol. 5, p. 216. Authors: Craig C. Freeman, James L. Reveal.
Parent taxa Polygonaceae > subfam. Eriogonoideae > Eriogonum > subg. Ganysma
Sibling taxa
E. abertianum, E. acaule, E. alatum, E. aliquantum, E. allenii, E. alpinum, E. ammophilum, E. ampullaceum, E. androsaceum, E. anemophilum, E. annuum, E. apiculatum, E. apricum, E. arborescens, E. arcuatum, E. aretioides, E. argillosum, E. argophyllum, E. arizonicum, E. artificis, E. atrorubens, E. baileyi, E. batemanii, E. bicolor, E. bifurcatum, E. brachyanthum, E. brachypodum, E. brandegeei, E. breedlovei, E. brevicaule, E. butterworthianum, E. caespitosum, E. capillare, E. cernuum, E. chrysops, E. cinereum, E. cithariforme, E. clavatum, E. clavellatum, E. codium, E. collinum, E. coloradense, E. compositum, E. concinnum, E. congdonii, E. contiguum, E. contortum, E. correllii, E. corymbosum, E. covilleanum, E. crocatum, E. cronquistii, E. crosbyae, E. cusickii, E. darrovii, E. dasyanthemum, E. davidsonii, E. deflexum, E. deserticola, E. desertorum, E. diatomaceum, E. diclinum, E. divaricatum, E. douglasii, E. eastwoodianum, E. effusum, E. elatum, E. elegans, E. elongatum, E. ephedroides, E. eremicola, E. eremicum, E. ericifolium, E. esmeraldense, E. evanidum, E. exaltatum, E. exilifolium, E. fasciculatum, E. flavum, E. fusiforme, E. giganteum, E. gilmanii, E. glandulosum, E. gordonii, E. gossypinum, E. gracile, E. gracilipes, E. gracillimum, E. grande, E. greggii, E. gypsophilum, E. havardii, E. heermannii, E. helichrysoides, E. hemipterum, E. heracleoides, E. hieracifolium, E. hirtellum, E. hirtiflorum, E. hoffmannii, E. holmgrenii, E. hookeri, E. howellianum, E. hylophilum, E. incanum, E. inerme, E. inflatum, E. intrafractum, E. jamesii, E. jonesii, E. kelloggii, E. kennedyi, E. kingii, E. lachnogynum, E. lancifolium, E. latens, E. latifolium, E. lemmonii, E. leptocladon, E. leptophyllum, E. libertini, E. lobbii, E. loganum, E. lonchophyllum, E. longifolium, E. luteolum, E. maculatum, E. mancum, E. marifolium, E. mensicola, E. microthecum, E. mitophyllum, E. mohavense, E. molestum, E. mortonianum, E. multiflorum, E. natum, E. nealleyi, E. nervulosum, E. nidularium, E. niveum, E. nortonii, E. novonudum, E. nudum, E. nummulare, E. nutans, E. ochrocephalum, E. ordii, E. ostlundii, E. ovalifolium, E. palmerianum, E. panamintense, E. panguicense, E. parishii, E. parvifolium, E. pauciflorum, E. pelinophilum, E. pendulum, E. pharnaceoides, E. plumatella, E. polycladon, E. polypodum, E. prattenianum, E. prociduum, E. pulchrum, E. pusillum, E. pyrolifolium, E. racemosum, E. reniforme, E. ripleyi, E. rixfordii, E. robustum, E. rosense, E. roseum, E. rotundifolium, E. rubricaule, E. rupinum, E. salicornioides, E. saxatile, E. scabrellum, E. scopulorum, E. shockleyi, E. siskiyouense, E. smithii, E. soliceps, E. soredium, E. spathulatum, E. spectabile, E. spergulinum, E. sphaerocephalum, E. strictum, E. subreniforme, E. suffruticosum, E. temblorense, E. tenellum, E. ternatum, E. terrenatum, E. thomasii, E. thompsoniae, E. thornei, E. thurberi, E. thymoides, E. tiehmii, E. tomentosum, E. trichopes, E. tripodum, E. truncatum, E. tumulosum, E. twisselmannii, E. umbellatum, E. ursinum, E. vestitum, E. villiflorum, E. vimineum, E. viridescens, E. viridulum, E. viscidulum, E. visheri, E. watsonii, E. wetherillii, E. wootonii, E. wrightii, E. zionis
Subordinate taxa
Polygonaceae subfam. Eriogonoideae, Polygonaceae subfam. Polygonoideae
Name authority Bentham: Trans. Linn. Soc. London 17: 406, plate 18, fig. 1. (1836) Jussieu
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