Flora of North America species comparison

Rumex acetosella in the broad sense is an extremely variable and taxonomically complicated polyploid complex, which includes diploids, tetraploids, hexaploids, and octoploids. This complex (excluding more distantly related arctic-montane R. graminifolius and its allies) probably originated and developed mostly in southern Europe and southwestern Asia. Some races of R. acetosella now are distributed almost worldwide as introduced and often completely naturalized aliens.

Á. Löve (1941, 1983) assumed that in this group chromosome numbers are strictly correlated with morphology. In his opinion, every chromosome race represents a distinct species: diploid Rumex angiocarpus Murbeck [= Acetosella angiocarpa (Murbeck) Á. Löve]; tetraploid R. multifidus Linnaeus [= R. tenuifolius (Wallroth) Á. Löve = Acetosella multifida (Linnaeus) Á. Löve]; hexaploid R. acetosella in the narrow sense [= A. vulgaris (W. D. J. Koch) Fourreau, with gymnocarpous A. vulgaris subsp. vulgaris and angiocarpous A. vulgaris subsp. pyrenaica (Pourret ex Lapeyrouse) Á. Löve]; hexaploid R. graminifolius Rudolph ex Lambert [= A. graminifolia (Rudolph ex Lambert) Á. Löve]. However, the distribution given by Löve for these taxa seems unnatural. Studies by J. C. M. den Nijs and collaborators (den Nijs 1974, 1976, 1984; den Nijs and T. Panhorst 1980; den Nijs et al. 1980, 1985; see also W. Harris 1969, 1973) indicate that the situation is more complicated. They postulated the development of two major evolutionary lines into two ploidy complexes: a primary western Mediterranean one and a secondary eastern Mediterranean one. According to this scheme, polyploid races independently and spontaneously emerged (and still are emerging) within different ancestral populations.

The most widespread, almost cosmopolitan race, presumably native to the southwestern Mediterranean region, including southwestern and Atlantic Europe, which is common in North America, is characterized by a hexaploid chromosome set (2n = 42), nonmultifid lateral lobes of basal leaves, and angiocarpy (fruits are not easily separable from accrescent inner tepals). It was commonly and erroneously referred to as Rumex angiocarpus Murbeck, or R. acetosella subsp. angiocarpus (Murbeck) Murbeck. According to J. R. Akeroyd (1991), who in general followed the taxonomic revision of the group by J. C. M. den Nijs (1984), the correct name for this taxon is R. acetosella subsp. pyrenaicus (Pourret ex Lapeyrouse) Akeroyd (=Acetosella vulgaris subsp. pyrenaica (Pourret ex Lapeyrouse) Á. Löve). Gymnocarpous nonmultifid and multifid forms (R. acetosella subsp. acetosella and R. acetosella subsp. acetoselloides (Balansa) den Nijs, respectively) also occur in North America, but evidently rarely. The distributions of subspecies of R. acetosella in North America are poorly known. Keys and detailed descriptions for the subspecies were provided by den Nijs and Akeroyd. However, the tempting simplicity of the keys is somewhat suspicious. The alternative point of view (and an alternative key) may be found in Á. Löve (1983).

Rumex acetosella subsp. arenicola Mäkinen ex Elven was recently described from Greenland and reported for Scandinavia and arctic Russia (R. Elven et al. 2000). This entity seems to be morphologically transitional toward Rumex graminifolius (see discussion under that species below). According to Elven et al., it differs from other infraspecific entities of R. acetosella in having the following characters: leaves usually without basal lobes (as in R. graminifolius), with revolute margins; inflorescence sparsely branched; tepals and pedicels densely covered with red papillae (as in R. graminifolius). From R. graminifolius and related taxa (R. beringensis and R. krausei) it can be distinguished by narrower inner tepals (similar in size to those in other subspecies of R. acetosella). The distribution of subsp. arenicola and its relations to other taxa are in need of further study.

(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.)