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AGAIN: TAXONOMY OF YELLOW-FLOWERED CAULESCENT OXALIS (OXALIDACEAE) IN EASTERN NORTH AMERICA PDF

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Preview AGAIN: TAXONOMY OF YELLOW-FLOWERED CAULESCENT OXALIS (OXALIDACEAE) IN EASTERN NORTH AMERICA

TAXONOMY AGAIN: OF YELLOW-FLOWERED CAULESCENT OXAUS NORTH (OXALIDACEAE) EASTERN AMERICA IN Guy L Nesom Yellow-flowered caulescent Oxalis (sect. Corniculatae) of North America has been the and object of interest more study than for century. Small number new a (1898, 1903, 1907) described and a of taxa brought the names wide he into usage; also treated sect. Corniculatae as separate genus, Wiegand a Xanthoxalis Small. (1925) provided a detailed and formal overview of the group and recognized many minor with variants formal names, with an emphasis on of study northern North America. Eiten's (1955, 1963) studies distilled taxonomy more the on to a set of taxa based realistic biological concepts rather than typological ones. Lourteig group and (1979) restudied the applied very names a different set of to the taxa, differing in part and in the selection of types in part in concepts of relationships and taxonomic ranks. Useful perspectives on problems and of typification nomenclature have been provided by Watson and Ward (1989) (2004), as noted below. problems significant in have remained and Still, identification confusion about which persists names are appropriate. In restudying, North American again, the eastern plants toward development of a treatment of Oxalidaceae for the Flora of North America have come series, to a different view, although I still than it is far closer to Eiten's Lourteig's. In fact, Lourteig's biology and nomenclature appears have been to many retrograde ways compared in to Eiten's. In the taxonomic summaries synonymy below, showing more commonly representative, the used is names and some that allude aspects of within synonymy to variability the species. Additional can be found in Eiten Lourteig and Watson USA (1955), (1979), (1989). Closely related taxa of the western are Oxalis call- & (Abrams) Knuth, fornica Kunth, R. 0. albicans O. pilosa Nutt. ex Torr. Gray, and Trel.— O. these suksdorfii of Texas Journal of the Botanical Research Institute 3(2) 28 7 Turner Wiegand 1994). Lourteig 1979; taxonomic arrangements 1925; have been in various (e.g., treated also Young and provided by (1958) Eiten (1963). Useful are illustrations with not or morphologically intergrading at all understood taxa little present species are as In the study, With further study, reproductive isolation. taxa— sympatry such taxa taken as evidence of their of other is show taxonomic probably the biology will reproductive complexities in beyond scope overview, the of this odd- and populations of framework advanced here be overly simplistic. In Oxalis corniculata 0. dillenii, to chromosome numbers have been reported; polyploid populations are and dysploid even-polyploid, polyploid, homostylous species corniculata, 0. study (1963) suggested that essentially (0. reported in 0. Eiten's stricta. when and pollen tube self-pollinated—pollination pseudogamous apomicts are O.florida, O. stricta) dillenii, Among be appears a facultative to endosperm these, at least O.florida growth only fertilization. resulting in when and producing sexual seeds abundant apomictic seeds also producing apomict— emasculated flowers may from outcrossing Sexual seed production Doust 1981). pollinated with acceptable pollen (Lovett et al. with morphological heterostyly 0. grandis, 0. priceae) Species strong (e.g., occur in the other species as well. combina- and hybrids various produced natural in hybrids are easily consistently outcrossers. Artificial are by have been Eiten (1963). identified tions Acknowl- 2400 specimens estimated (see on examination conservatively The study based of a present is among many specimens those studied and annotated Both Lourteig Eiten edgements herbaria consulted). for with concepts of their GH, SMU, and TEX-LL, so the current author is familiar in detail especially here, at Arkansas, Corniculatae in populations of author has studied natural sect. In 2009, the the taxa involved. Oklahoma, and Texas. Massachusetts, Missouri, Louisiana, NORTH AMERICA OF EASTERN YELLOW- FLOWERED CAULESCENT OXALIS SPECIES TO THE KEY and peduncles Stems evenly strigose from base to pedicels. 1 . mm complete 5-1 cymes, homostylous; petals long, 2-3(-5 umbelliform to Flowers or rarely 8) in 1 2. 1 ; with yellow, cym< umbelliform (2-)3-5(-8) in •s glabrous, Stems to to nearly rare pilose villous , mm promir 0-20 yellow with long, Petals 3. 1 Stems densely and pilose with sp stiffly 4. and an plant lignescent or individual lie prominently red-line of the leaves; corolla throats level densely or Stems glabrous to sparsely or pilose \ nearly 4. rhizomes usually and nonseptate stoloniform hairs; 1 above cymes, or within tr 2-4(-8) regular or irregular in mm 0-1 to strongly red-lined within; petals 8 Ic faintly 1 stoloniform from lignescent, slender, Plants arising 5. nan rounded, margins often with a shoulders usually mm 10-14 throaty long, the petals of leaves; level Petals4-9(-11)mmlong,y< 3. corn Oxalis and aur 1. free distal the an rooting at from the leaning singly bast rarely usually ing Stems' erect, aris 6. ;, obsolescent pules margins without or* auricles. free distal very Stems 20-60(-90) long, sparsely to err s 7. i fron singly densely arising or villou hairs ite ;, were (3-)5--7(-15) regular usually in ( Oxalis 6. jstopuberulentandvilk 729 Nesom, Taxonomy of yellow-flowered caulescent Oxalis Stems (5-)8-30(-35) cm, sparsely pilose with nonseptate hairs to almost completely glabrous, aris- 7. umbel ing from a taproot, often producing lignescent stolons; flowers or2(-3, rarely 4-5) in liform 1 cymes; capsules glabrous to sparsely puberulent, not villous 5. Oxalis florida (Thunb.) Zucc, Denkschr. Koeinj var. repens .lata Stems and Plants perennial, caulescent, arising from thin to lignescent taproots. 4-10(-30) cm, several radiat- ing laterally from the taproot, creeping, rooting at the nodes and stolonlike, prostrate to distally ascending- and Stipules herbaceous but lignescent, sparsely loosely strigose to strigose-villous. present, erect, initially Leaves and membranous, margins with basal cauline; obcordate, free flanges, distal auricles free. leaflets 3, mm, on maroon, margins lobed 1/5-1/3 length, (4-)6-12 green both surfaces or bronze-purple to often Flowers 2-3 umbelliform cymes, mostly prominently 1-5 cm. or (-6) in irregular or villous-ciliate, petioles 1 mm, 4-8 homostylous; peduncles (l-)2-4(-8) cm; pedicels in fruit horizontal to deflexed; petals yellow. mm, 8-17(-20) puberulent Capsules angular-columnar, gradually or abruptly tapering apex, sparsely to to glabrate or glabrous. Seeds uniformly brown, transverse ridges brown. In = 24, 36, 42, 44, 48. Flowering Mar-Aug, sporadically year in Florida. Gardens, greenhouses, lawns, fields, roadsides, all m; hammocks, beach margins, open pine woods, grasslands; 10-500(-2500) Nfld.!, Ont.!, P.E.I.!; Ala.!, Kay Ark.!, Colo.!, Conn.!, D.C.!, Ha.!, Ind. (fide Yatskievych), La.!, Me.!, Mass.!, Mo.!, N.C.!, Calif.!, N.J.!, 111.!, West America, South WYa.!; introduced; native Mexico, Indies, Central to Oreg.!, Pa.!, S.C.!, Tex.!, Vt.!, Va.!, PLANTS The America; introduced Europe, Asia (India, China, Japan), Africa, Pacific Islands, Australia. and Wash. Database shows records for Ariz., Ga., Nebr., Ohio, Okla, S.Dak., Many broad sense have been described over cosmopoli- infraspecific taxa of Oxalis corniculata in the its The and but taxonomic uncertain only the single entity treated here. reported tan range, their status is is concomitant complex morphological variation in ploidy level a of the variation. is USA recognized by small flowers, sparsely hairy stems creeping Oxalis corniculata in the relatively is its procumbent and from and developed with and rooting nodes, radiating the taproot, well- stipules its at all and Peduncles and 2-3 produced the nodes, broad, marginal flanges auricled apices. or leaves are at free 1 become through short erect stems rarely. Plants flower as annuals but often short-lived perennials the colonial may Stems sensu be decumbent or prostrate and rooting the nodes, but they habit. of O. stricto at dillenii and from stems charac- almost always are ligneous lignescent, not evidently radiating the taproot, erect to form teristically arise from the nodes. According to Eiten (1963), O. dillenii is able to "vigorous, floriferous, with but hybrids 0. corniculata." sterile maroon and pubescent have been with bronze-purple leaves capsules Plants of Oxalis corniculata to Ward Abrams Such recognized atropurpurea in Florida, 2004; in California, 1951). as O. corniculata war. (e.g., and whether popu- plants apparently occur sympatrically with the typical expression is not clear they are it lational variants or whether they are at least partially reproductively isolated. In Malaysia, var. atropurpurea and morphology from typical 0. corniculata in karyotype as well as in floral vegetative differs & 2004)— Kuriachan by post-pollination reproductive barriers (Nair at least in that regioi Aus behavior atropurpurea should be treated specific rank. Distinctive indicates that var. at 730 Journal of the Botanical Research Institute of Texas 3(2) ants sometimes identified as 0. corniculata have recently been treated as distinct species de Lange et (e.g., A common 2005). form of O. corniculata in central Mexico (collections seen from Hidalgo, Jalisco, Edo. is al. — Mexico, Michoacan, Morelos, and Queretaro) these plants produce large, prominent stipules and nearly glabrous stems, but the habit varies from procumbent to ascending and the stems rarely root at the nodes. Most American Americas and occur of the closest relatives of Oxalis corniculata (in expression) in the its USA, West Indies and the species probably native there, though probably south of the United States. In the is occurs mostly urban and highly disturbed but along the Gulf coast occasion- O. corniculata in habitats, it ally grows in less obviously disturbed sites. North of the Gulf Coast, its occurrence is mostly restricted to greenhouses and horticultural (lawns and gardens), suggesting that repeatedly introduced rather sites is it than persisting (and expanding) through seed production. Oxalis Mon. 28 N.Ame dillenii Jacq Oxalis 1794. d iA.GraySyn.Fl.: 1. Oxalis conMculatc L. var. Jacq. Trel. ir ) , . i 1(1):36: entirely se ated pre made Election by Marl* IDillenius :eivednea specimens fror ;by's merica (Reveal 1983). I 955, 104) observed "Although plate lsoapla p. lat i tl 1 herbariiun which is almost sure ly the plant Di edCDruce,SrVine s 1907, p. 176), and therefore:,ifunique,istheh olotype Zuc Amer. 1811 L (Pursh) Sept. 1:322. [1813]. O;cafe corniculata var. lyonii csh, Fl. ' .. 1800-1813], John Lyo lbyhta (asnotedbyEite, ange O.texan for U.var.^ridcmaWieg.,Rhodor 27:138. >25. Type: U.S.A. Florida: East Florida, rtodate, D.C. Eaton s.n 1! £ . ItaUSANsv. aWieg.,Rhodora27: C taocar 123. 1925. Hampshiei. [Cheshire :o.I:Als tead, old grave P P - :. Plants perennial, caulescent, arising from a ligneous or lignescent rhizome, sometimes appearing taproot- mm 2-8 Stems 10-25 cm, proximally ligneous lignescent, 1.5-2 thick, usually from the base, erect like. to becoming decument and sometimes initially, often to prostrate rhizome-like, rooting at nodes, strigillose to strigose with antrorsely appressed, nonseptate, sharp-pointed hairs. Stipules usually greatly reduced at above midstem, margins narrowly without any without least flanged or free portion, free apical auricles. mm, Leaves and lobed 1/5-1/3 (4-)6-15(-21) green on both basal cauline; obcordate, length, leaflets 3, 1-4 Flowers cm. 2-3(-4, very surfaces, glabrous adaxially, sparsely strigillose abaxially, petioles 1 or rarely cm; umbelliform cymes, mostly homostylous; peduncles l-6(-10) pedicels in to 8) in (rarely irregular) fruit mm, 5-11 Capsules deflexed horizontal), usually without bracteoles; petals yellow. angular-columnar, (to mm, and abruptly tapered apex, 12-20(-25) densely strigose-pilose with mixture of appressed spreading at with puberulent understory. Seeds brownish, tranverse ridges with strong grayish or white hairs, a lines. In = 20-24. 20, 22, 18, Flowering Feb-May(-Oct). Pastures, roadsides, lawns, river bottoms, sandy, rocky, or gravelly soils; . Nesom, Taxonomy of yellow-flowered caulescent Oxalis 731 5-300 m; N.B.!, N.S.!, Ont.!, P.E.I.!, Que.!, Sask.!; Ala.!, Ark.!, Ariz.!, Colo.!, Conn.!, Del!, D.C.!, Fla.!, Ga.!, Ind.!, Iowa!, Kans.!, Ky.!, La.!, Maine!, Md.l, Mass.!, Mich.!, Minn.!, Miss.!, Mo.!, Nebr.!, N.H.!, 111.!, N.J.!, N.Mex.!, N.Y.I, N.C.!, N.Dak.!, Ohio!, Okla.!, Pa.!, R.I.!, S.C.!, S.Dak.!, Tenn.!, Tex.!, Va.!, Vt.!, Wash.!, W.Va.!, PLANTS Wis.!, Wyo.!; introduced in Bermuda, Europe. The Database shows records from Man., Idaho, Alta, and Oreg. Decumbent branches appear producing branches and of Oxalis often as stoloniform, erect leaves dillenii at the nodes, rarely producing a few, small adventitious roots. Such plants sometimes are misidentified as 0. and corniculata, but they differ in their overall habit, ligneous stems rhizomes, reduced stipules, strigillose and cauline vestiture, denser fruit vestiture, seed color. In 0. the stems are consistently strigose, dillenii, commonly while the long petioles are pilose-villous, and in a quick scan of a specimen not carefully pressed, may continue begin moist Oxalis to flower, or to reflower, in places especially in the Plants dillenii fall. mowed commonly abundant lawns and and are often in other areas, later-flowering plants produce very much may The shows some which small be flowers. species plasticity in habit, of genetically partitioned, as chromosome dysploid races apparently exist. Oxalis and O. stricta are the two most abundant species of the eastern United States and both dillenii commonly comments occur in disturbed In view of recent nomenclatural confusion below, under (see sites. 0. a key that separates these two presented here. stricta), is Stems usually 2-8 from the base, 0-25 cm, erect often becoming decumentto prostrate and rhizome- initially, 1 1 cymes or lignescent rhizomes or stolons; flowers or 2-3(-5, rarely to 8) in umbelliform Oxalis dillenii 1 2-3 Stems usually arising singly from the base (rarely together), 20-60(-90) cm, erect or later leaning or fall- 1 . ing over and decumbent; stem vestiture very sparsely to sparsely or moderately pilose or villous with non- Club Torrey 25:612. 1898. Bull. Bot. Xanthoxdis priceae (Small) Small, S.E. U.S. 668. 1903. Fl. ^Y, NY-digital image!). Plants from with perennial, caulescent, arising a ligneous or lignescent taproot, usually lignescent, stoloni- form (or offset-like) rhizomes rooting at nodes and producing erect stems from the nodes. Stems proximally 2-8 from 5-20(-40) cm, becoming decumbent, lignescent, usually the base, erect or usually villous-hirsute with nonseptate hairs spreading or deflexed or spreading in dissimilar orientations. Stipules with margins narrowly without any without Leaves and flanged or free portion, free apical auricles. basal cauline; leaflets 3, mm, obcordate, lobed 1/5-1/3 length, 3.5-12 green on both surfaces, strigose-hirsute on both surfaces, less commonly glabrate, petioles 2-7 cm. Flowers or (2-)3-8 in umbelliform cymes, less commonly irregular 1 cymes, produced well above the level of the leaves, distylous; peduncles (3-)5-10(-15) cm; pedicels in fruit Journal of the Botanical Research Institute of Texas 3(2) 732 mm, prominent (13-)14-20 with red the deflexed horizontal, often bracteolate; petals yellow, stripes at to mm, 10-15 Capsules abruptly tapered apex, sparsely to densely base angular-cylindric, at (corolla throat). commonly puberulent with deflexed with nonseptate short, straight, hairs. villous long, deflexed, hairs, less unknown. In = Seeds usually with white transverse ridges. and Flowering Mar-May. Dry limestone areas, glades, cedar barrens, chalk prairies, limestone bluffs 5-300 m; sandstone rocky sandy hedgerows, oak-pine, longleaf pine; Ala.!, outcrops, slopes, talus, cliffs, Mexico (Nuevo N.CJ, Tenn.!; Leon!). Ha.!, Ga.!, Ky.!, Miss.!, S.C.?, USA, by recognized vinous-hirsute species of the southeastern villous to Oxalis priceae a distinctive its is umbelliform cymes, and yellow yellow-orange corollas with red lines in the throat. stems, flowers in large to The remain drying and usually can be seen on herbarium specimens even in the throat visible after lines A and from the outside of the flower. similar pattern also occurs in 0. texana, 0. illinoensis, 0. grandis. Plants of Oxalis priceae in Mississippi and eastern Louisiana grow in pineland and have slightly smaller corollas than those eastward. Vouchers for Lousiana records are cited here. Jefferson Par.: at Gretna, opposite RR Zimmerman New May Red near frequent Orleans, 6 1899, Ball 352 (GH); Rapides Par.: River valley, Sta., Tammany mi on 24 Apr Ewan 17608 (MO); in dryer spots beneath pines crest of hills, 1948, St. Par.: 5 tall US hwy embankment, 30 Apr Ewan 18546 (GH). 1953, S of Pearl River, 90, Two collections from a single area in North Carolina are typical Oxalis priceae, at the northeastern Apr extremity of main range. Stanly Co.: rocky slopes below bluffs of Yadkin River, near Charlotte, 20 its power 1932, Palmer 39985 (GH); steep moist banks above the Yadkin River, just above the second or lower dam May from 6826 eastern east of Badin, 10 1963, Wilbur (GH). Similarly, a collection of Oxalis priceae Newport Apr Tennessee appears be Cocke near Del Rio, siliceous bluffs along Rd., 18 1963, disjunct. Co.: to may known (TENN Sharp 17255 Further study in the distribution, especially in digital image!). et fill at. no Georgia and South Carolina, so that the Cocke Co. and Stanly Co. localities longer appear disjunct. from Two from montane Nuevo Leon, Mexico, are unmistakably Oxalis priceae, disjunct collections USA southeastern by about 900 kilometers. Nuevo Leon. Mpio. Santiago, along the closest localities in the the Cola de Caballo-Laguna Sanchez road into high sierra SE of Monterrey, between Puerto Genovevo and NW km dominated by La Puerto Genovevo, temperate pine-oak-hickory Pinus, Cienega, 1.5-2.5 of air ca. 1500 m, Mar 1887 (TEX); Mpio. Bustamante, along switchbacks of road be- Carya ovata, 15 1994, Mayfield km below and from opening 4 road low the Grutas de Bustamante, ravines bluffs along road of cave to ca. 1000 m, rocky Quercus-Cercis-Brahea-Ungnadia-Fraxinusgreggii, dark, rich, soil in talus slopes, fairly cave, common Mar E 1908 (TEX). in moist ravine of cave opening, 18 1994, Mayfield to Amer. xana (Small) Eiten, Midi. woody from caudex Plants perennial, caulescent, arising from a taproot, caespitose with stems arising a or Stems 5-15 with rhizome-like or stolons rooting the nodes. cm, proximally ligneous to lignescent, offsets at with appressed ascending nonseptate Stipules ascending, antrorsely to hairs. erect to strigose to strigillose rounded and Leaves usually with margins very narrowly flanged, usually with slightly free apical auricles. mm and lobed 1/5-1/3 the length, (4-)6-12(-18) long, green to purple basal cauline, cordate, leaflets 3, cm 2-6 on both surfaces, glabrous to sparsely strigose adaxially, sparsely strigose abaxially; petioles long. 4-10 Flowers (2-)3-5(-8) in umbelliform cymes, very rarely irregular cymes, distylous, peduncles cm; mm with without (10-)12-16(-17) long, yellow, pedicels horizontal deflexed in bracteoles; petals to fruit, Capsules angular-columnar, abruptly tapered apex, prominent red lines the base (corolla throat). at at mm 8-15 moderately densely puberulent puberulent-villous. Seeds brownish, transverse ridges long, to to unknown. In = whitish. distinctly Commonly Mar-May and but Flowering in undisturbed habitats usually in deep, loose sand, (-Jun). and mixed hardwoods; 10-200 m; and edges and openings woods, pine, pine-oak, roadsides, also fields, Ark.!, Tex.! La.!, Nesom, Taxonomy of yellow-flowered caulescent Oxalis 733 more numerous from Oxalis texana very similar to 0. dillenii— differing O. primarily in is dillenii its and The flowers per inflorescence distylous flowers with red-lined corolla throats. distinctive red larger, striping in the corolla throat remains visible after drying and usually can be seen on herbarium specimens even from and the outside of the flower. Plants of O. texana also are distinct in their relatively larger taproots on with with habit either caespitose or short stolon-like offsets. Plants of 0. larger flowers elevated dillenii peduncles might be mistaken for O. texana, yet the two taxa exist sympatrically in the range of 0. texana and seems and geography and clear that they are separate species. Oxalis priceae O. texana are separate in it morphology. name Lourteig (1979) used the Oxalis Pursh for the species identified here as 0. texana, and her lyonii drawn illustrations of 0. lyonii were from a Texas collection identified here as O. texana (Lourteig 1979, Fig. mapped and same Turner (2003) followed Lourteig's nomenclature the species as 0. a-e). et al. lyonii. 7, and Lourteig neotypified O. lyonii to place within her concept of O. texana (as identified here) included it — synonymy 0. [Xanthoxalis] texana in her neotype, however, from southern Mississippi, outside of the is him range of O. texana, and is instead a collection of 0. dillenii (as noted by Eiten 1963 and annotated by in comments synonymy 1965; confirmed here as 0. Further regarding O. are in the of O. dillenii). lyonii dillenii in the present paper. NY, NY-digital image!). itpe: jlorida Salisb. var. jilipes (Small) Ahles, Elisha Mitchell Sci. lis J. 57. 1903. Oxalis Jacq. subsp. (Small) Eiten, Amer. dillenii filipes North Yadkin 100-200 23 Aug 1894 J.K 59:301. 1963. Type: U.S.A. Carolina. Stanley Co.: Falls of the River, ft, Eiten 1963: NY, NY-digital image!; duplicates o pe, v etch Bot. S.C. 1:526. 1821. Oxalis jlorida Salisb. subsp. recurva C.E Reed, Phytologia 63:4 (Ell.) }. ARL, photo-GH!). Eiten (1963) notec would Geography e that place with equal probability as ei her 0. priceae [subsp. priceae] or 0. jlorida. places it Plants from youngest from perennial, caulescent, arising slender lignescent stolons, plants short, slender taproots. Stems usually single from the base, (5-)8-30(-35) cm, erect or rarely leaning and decumbent, sometimes strigose distally or just beneath the flowers and pilose-villous on proximal portions. Stipules obsolescent, without free margins or apical auricles. Leaves basal and cauline, leaflets obcordate, lobed 3, mm 1/5-1/3 length, 4-11 long, green on both surfaces, sparsely strigose abaxially; petioles 2-5 cm. Flow- ers 1 or 2(-3, rarely 4-6) in umbelliform cymes at level of the leaves or slightly above, tristylous; peduncles mm, Capsules (2-)3-8 cm; pedicels reflexing ascending, often bracteolate; petals 5-9(-ll) yellow. to mm, Seeds angular-cylindric, 8-12(-15) glabrous sparsely puberulent. brownish, including transverse to In = ridges. 16. Low swamp Flowering Mar-May(-Aug). woods, woods, pine woods, sandy burned- rich forests, sites, over woods, ditches, roadside banks, floodplains, low lake edges, creek banks, pastures, disturbed fields, 10-350 m; Kay rocky sites, bluffs, slopes; Ala.!, Ark.!, Conn.!, D.C.!, Fla.!, Ga.!, Ind. (fide Yatskievych), Ky.!, Maine!, Md.!, Mass.!, Mo.!, N.C.!, Tenn.!, W.Va.! La.!, Miss.!, NJ.!, N.Y.!, Pa.!, S.C.!, Vt.!, Va.!, 734 Journal of the Botanical Research Institute of Texas 3(2) and without recognized by mostly stems, sparse spreading cauline vestiture Oxalisflorida erect is its multicellular hairs, obsolescent stipules, relatively small, and yellow flowers without red lines in the throat; the stems and peduncles are thin compared to other species. a species primarily of the Atlantic distal It is known much and Gulf more sparsely represented in more inland regions. from a cluster of states coast, It is Wayne and MO!), and from southern Arkansas Missouri counties in southeastern (Butler, Carter, Ripley, cos.; (Ashley, Bradley, and Ouachita BRIT!, NLU!) northward to apparently isolated localities in Arkansas cos.; waif— The voucher West record perhaps was Tucker Otter Creek MO!). the Virginia a Co.: Yell Co.; for (e.g., A Lumber Co., near Hendriks, dry grounds along railroad track, 10 Sep 1904, Greenman 400 (GH). collection annotated by Eiten interpreted here as O. with slightly reduced vestiture, as 0. dillenii var. is dillenii filipes damaged Apr Tharp perhaps resulting from stems: Polk Co: 12 1941, (GH). s.n. Ward and noted Lourteig (1979) also treated Oxalisflorida at specific rank, for Florida, (2004, p. 35) and between and appreciable that "the differences O. subsp. subsp. (= O. florida) "are dillenii dillenii filipes intermediates seem few." In contrast, Eiten (1963, 268) observed that Oxalis dillenii subsp. filipes "is dis- p. tinctive in its most characteristic form but intergrades with [subsp. dillenii], both in forming intermediate homogeneous populations and in disturbed ground, variable hybrid swarms. The most distinctive por- also, The northern Appalachians. morphological evidence tion of this variable subspecies concentrated in the is from whether from conflicting as to originated [subsp. dillenii] or directly [O.] corniculata." is it Wiegand preceding Eiten, also observed a measure of intermediacy in Oxalisflorida, (1925, 124), p. much between and noting that "0. and O. have the appearance of hybrids [0. stricta either 0. florida filipes or 0. corniculata], as no new characters are found in either species. The frequency of their occurrence dillenii in the east and absence in the west, where the possible parents both occur is against this hypothesis." name Ward was noted Eiten incorrect in claiming that the Oxalisflorida Salisb. (2004, 35) that is p. name synonymy by was pre-Linnaean which, "The [phrase-name] cited in Salisbury (1796) illegitimate. since not available for his use, does not disturb the legitimacy of O. florida" and was In view the difference between Oxalis subsp. O. priceae subsp. colorea Eiten's (1963), dillenii filipes mm keyed primarily in flower Subsp./zlipes keyed under "Flowers 13 long or less," while subsp. colorea size. mm under "Flowers 10 long or more." The seemingly arbitrary difference in size apparently is reflected in comment his that he identified a duplicate (F) of the type of Xanthoxalis colorea as O. dillenii subsp. filipes. mostly In the current study, have sorted collections previously identified as Oxalis priceae subsp. colorea I between and and the of subsp. colorea remains somewhat in doubt, even though O. O. priceae, reality florida synonym Mulcahy found both sensu placed here as a of 0. priceae. (1964, 1048) that O. priceae stricto is p. it and 0. priceae subsp. colorea exhibit a high degree of self-fertility, the former strictly distylous, the latter and between forms very dense "A further difference the 2 subspecies that subspecies priceae tristylous. is extensive clones, some of which extend for several meters, while subspecies colorea forms rather diminutive 3-4 Ward very few which contain more than flowering did not find significant variation clones, of stalks." synonym. within O. priceae, listing Xanthoxalis colorea as a 6. Oxalis Stricta L, Sp. Pi. 1:435. 1753. Type: Morison, PL Hist. Univ. 2:184, 2, 17, fig. 3. 1680 Optotype. Eiten 1955). s. t. The protologue noted "Habitat in Virginia." iii(Small)H.Hara,J.Jap.B Nesom, Taxonomy of yellow-flowered caulescent Oxalis ima ellBOTYEKN 1 m, 2 1900, PA. Rydberg 5920 with F K. Vreeland (holotype NY, NY-digital NY-digital image!). Jul if, Rhodora cymosa Torrey Club (Small) Wieg., 27:135. 1925. Type: Small, Bull. Bot. ioscl alis New U.S.A. York. [Bronx Co.]: Riverdal NY-digital image!). Appar ntlynotypee .1854. Type: Wiegand noted "0. uropam with d DubtanativeofAme ntroduced into Europe about 1658. (1927, 136) that is little p. ner 7:149]." [1913, Vol. May C 20 mropaea Jordan forma pilosella Wieg. Rhodora 27:135. 192 5.Type:U.S.A.Missouri. Jackson o.: Courtney [Greenwood], alis Stems Plants annual short-lived perennial, caulescent, arising from a thin, short rhizome. usually arising to 2-3 and decumbent, 20-60(-90) from leaning or over singly the base (rarely together), erect or later falling commonly on and concen- stems cm, withnonseptate spreading septate hairs present petioles, villous hairs, nodes, sometimes only on Stipules obsolescent, without free margins or apical auricles. trated petioles. at mm, green Leaves and obcordate, lobed 1/5-1/3 length, (8-)10-20(-30) light to basal cauline; leaflets 3, 2-8 Flowers cymes, on both cm. usually (3-)5-7(-15) in regular yellowish-green surfaces, petioles rarely 1 commonly homostylous in irregular cymes, usually within level of the leaves or slightly above, or slightly less strongly heterostylous; peduncles 3-9(-ll) cm; pedicels in fruit erect to ascending, often bracteolate; to mm, mm, 8-15 (6-)8-ll Capsules columnar, nearly abruptly tapering toward apex, petals yellow. terete, Seeds brown, In = with transverse ridges rarely whitish. 24. villous septate hairs to glabrate. 18, Flowering (Apr-)Jul-Oct. Prairie ravines, river and stream banks, sand bars, low woods, floodplains, 20-1200 m; roadsides, lawns, gardens; B.C., Man., N.B.!, Nfld., N.S.!, Ont.!, P.E.I.!, Que.!; Ala.!, Ark.!, fields, Colo.!, Conn.!, Del.!, D.C.!, Ga.!, Idaho, Ind.!, Iowa!, Kans.!, Ky.!, La.!, Maine!, Mass.!, Md.!, Mich.!, 111.!, Minn.!, Miss.!, Mo.!, Nebr.!, N.H.!, NJ.!, N.Y.!, N.C.!, N.Dak.!, Ohio!, Okla.!, Pa.!, R.L!, S.C.!, S.Dak.!, Tenn.!, (New The W.Va.!, Wis.!; introduced in Europe, Asia, Africa, Pacific Islands Zealand), Australia. Va.!, Vt.!, PLANTS and Wyo. shows from Mont., N.Mex., Wash., Database records Nfld., Sask., Ariz., recognized by erect stems from a short, simple rhizome, presence of septate Oxalis stricta is its tall, on and cymose and small Septate hairs the stems petioles are easily inflorescence, relatively flowers. hairs, do recognized especially because of their brownish crosswalls, but they vary greatly in density, as the (lens), dense and nonseptate hairs. In "villicaulis" and "pilosella" forms, as well as "var." bushii, the septate hairs are more around nodes and intermixed with evenly distributed on the stems, but often they are localized the nonseptate Often they are few in number, and, in rare cases, plants with greatly reduced vestiture hairs. overall apparently lack septate hairs. and and The rhizomatous lacking taproots, of Oxalis close relatives O. grandis O. illinoensis habit, stricta its The incompletely distinctive and a basic biological difference. bases of Oxalis plants often are collected, is however, especially in the other species, which commonly develop stoloniform or rhizomiform branches above the taproot. uncommon and from Texas western probably adventive. Reports of the species Oxalis in states stricta is 736 Journal of the Botanical Research Institute of Texas 3(2) NRCS USDA, 2009) have not been confirmed in the present study and unlikely that such reports is (e.g., it are accurate. The early establishment of O. stricta in Europe is noted above with the type of 0. europaea. may North Eiten (1963) appeared to suggest that the species be native in eastern Asia as well as America. comments 304-305) on and "Very in the species are insightful repeated here: Eiten's (1963, variability p. enough both between Asia and North America, and within each region. have not seen variable, as east 1 up North America material from Asia determine useful subspecies. In the variation east to is to set all if it up The most based regionally too intergrading useful infraspecific categories. distinctive that to set is is geographically-based character the presence of hairs on the upper surface of the This most is leaflets. is and frequent in the Midwest, but even there less than half the collections in each state have this feature, up and have been used sometimes varies even within single populations. This other characters that to set it same infraspecific groups (presence of septate hairs on stems or on pedicels) often vary in the population or even on the same plant, and present intergrades of expression. In addition, the extreme expressions all The vary they do not vary introduced of characters that regionally center in different areas, together. i.e., European North American being pubescent. plants represent a small part of the variation, in general less They not are at like the plants of east Asia." all Us (Small) Small, S.E. U.S. 668. 1903. Type: Fl. Stems from rhizomes without Plants perennial, caulescent, arising slender, lignescent, stoloniform tubers. cm 2-3 usually single from the base (rarely together), (10-)25-60(-100) before branching, nearly erect, with mixture and nonseptate Stipules glabrous to sparsely or densely pilose or villous a of septate hairs. 5-25 absent. Leaves cauline, mostly on the distal half of the stem; leaflets obcordate, lobed 1/5 length, (-30) 3, mm, on both margins sometimes upper rounded, green shoulders of lobes usually rarely flattened, surfaces, cymes Flowers 2-4(-8) narrowly brownish-purple, 5-7.5 cm. or in regular or irregular or petioles ciliate, 1 umbelliform cymes produced above the level of the leaves, tristylous; peduncles 7-12 cm; pedicels in fruit mm, 10-14 weakly Capsules erect to ascending; petals yellow, throat yellow to faintly or red-lined within. mm & 6-10 Seeds brown, In = 28 Weller ovoid to ovoid-oblong, long. traverse ridges rarely whitish. (fide Denton 1976, reporting unpublished counts by Ornduff). May-Aug. Sandy woods and 100-1100 m. Flowering Md.!, alluvial soils; Ala.!, D.C.!, Ga.l, Ind.!, Ky.!, WYa.! N.C.!, Ohio.!, Penn.!, S.C.!, Tenn.!, Va.!, & A report of Oxalis grandis from southern Mississippi (Forrest Co.; Carter Jones 1968) far out of is on which and and probably based perhaps range habitat for the species a misidentification, of O. priceae, is also large-flowered. is from herbaceous rhizomes producing white Plants perennial, caulescent, arising slender, at intervals fusi- 15-40 Stems 2-3 form tubers or tuberlike thickenings. usually single from the base (rarely together), erect, and cm, nearly glabrous or sparsely densely villous with a mixture of septate nonseptate hairs. Stipules to Leaves obsolescent, without free margins or apical auricles. cauline; leaflets obcordate, lobed 1/5 length, 3, mm, (12-)20-30(-35) upper shoulders of lobes green on both surfaces, margins green, petioles 4-7.5 ciliate, flat, cm long. Flowers l-3(-6) in regular or irregular cymes produced mostly at the level of the leaves, tristylous; mm, 3-10 12-18 peduncles cm, pedicels in fruit erect to ascending; petals yellow, throat strongly red-lined mm. Capsules 7-10 Seeds brown, In = unknown. within. oblong-ovoid, traverse ridges rarely whitish.

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