chorizanthe, spineflower
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Parry's spineflower, San Bernardino spineflower, San Fernando Valley spineflower
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Herbs [or subshrubs], annual [or perennial]; taproot slender to stout. |
Plants prostrate to spreading, 0.2–0.8(–1) × 0.5–4(–6) dm, strigose. |
prostrate or decumbent to erect, pubescent; aerial flowering stems arising [at nodes of caudex branches, at distal nodes of aerial stems or] directly from the root, decumbent to erect, slender [to stout and solid, not disarticulating in ringlike segments], sometimes disarticulating at each node. |
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persistent or quickly deciduous, basal and rosulate or basal and cauline, alternate; petiole present; blade linear to oblanceolate or spatulate, entire apically. |
basal; petiole 0.5–2(–3.5) cm; blade oblanceolate to narrowly oblong, 0.5–2.5(–4) × 0.2–0.6(–1.2) cm, thinly pubescent. |
terminal, cymose or capitate, uniparous due to suppression of secondaries; branches open and spreading or erect, typically trichotomously branched at proximal node, otherwise dichotomous, sometimes brittle and disarticulating into segments, round, pubescent [or rarely glabrous]; bracts mostly 2, opposite, sometimes numerous, whorled, distinct, leaflike to subulate or linear, occasionally awn-tipped, thinly pubescent (sometimes appressed), hirsute, villous, strigose, or tomentose, rarely woolly-floccose or minutely glandular. |
with involucres in small, open clusters 0.3–1 cm diam., greenish or grayish to reddish; bracts 2, sessile, usually leaflike, oblanceolate to elliptic, 0.5–1.5 cm × 1.5–7 mm, gradually reduced and becoming scalelike at distal nodes, linear, aciculate, acerose, 0.1–0.5 cm × 1–2 mm, awns straight, 0.4–1 mm. |
absent. |
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1–6+ per node, 3–6-ribbed, tubular, cylindric to urceolate or turbinate to campanulate; teeth 3, 5, or 6, awn-tipped. |
3–5, greenish or grayish to reddish, urceolate, slightly ventricose basally, 1.5–2 mm, corrugate, without scarious or membranous margins, pubescent; teeth widely spreading to divergent or recurved, equal, 0.5–1.5 mm or 1–3 mm; awns uncinate or straight, unequal, alternating 0.5–1.5 mm and 0.2–0.5 mm. |
bisexual, 1(–2) per involucre, pedicellate; perianth white to yellow or pink to rose-pink, red, maroon or purple, cylindric, funnelform, or campanulate when open, cylindric when closed, glabrous or pubescent abaxially; tepals (5–)6, connate 2/3 their length, monomorphic or dimorphic, entire, emarginate, or lobed to laciniate apically; stamens 3, 6, or 9, or variously 3–9; filaments distinct or connate into staminal tube, sometimes adnate to floral tube, glabrous or pubescent; anthers maroon to red or cream to white or yellow, oval to oblong; styles erect to spreading. |
slightly exserted; perianth bicolored with floral tube greenish white and tepals white, cylindric, 2.5–3 mm, sparsely pubescent; tepals connate 2/3 their length, slightly dimorphic, those of outer whorl oblong to oblong-ovate, 1.5 times longer than those of inner whorl, rounded, erose or rarely some entire to denticulate apically, those of inner whorl linear-oblanceolate, acute, entire or denticulate apically; stamens 9, included; filaments distinct, 2–2.5 mm, glabrous; anthers white, ovate, 0.2–0.3 mm. |
included, light brown to dark brown or black, not winged, lenticular, globose-lenticular, or 3-gonous, glabrous. |
brown, globose-lenticular, 2.5–3 mm. |
embryo straight or curved. |
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= 10. |
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South America; North America (including Mexico) |
CA
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Species 50 (33 in the flora). Like Eriogonum, Chorizanthe is the basal element in its own subtribe, Chorizanthineae Reveal. Nonetheless, recent molecular data indicate that Chorizanthe is embedded within Eriogonum (A. S. Lamb Frye, pers. comm.), meaning that either all species of the Chorizanthineae should be moved to Eriogonum, or Eriogonum should be fragmented into several genera. Obviously, therefore, all of the segregate genera that follow could be merged into Chorizanthe, and this was common practice until 1989. A key factor still unresolved is the relationship between the perennial species of Chorizanthe (including the type of the genus) and Eriogonum. The traditional assumption is that Eriogonum and Chorizanthe represent independent lines of evolution from a basal, diploid (n = 10) ancestor that is now extinct. A corollary to this assumption is that this divergence occurred during the Eocene and the perennial spineflowers were successfully introduced into southern South America, whereas the wild buckwheats—lacking a ready means for long-distance dispersal—failed to make the trip. Thus, it is possible that the perennial members of Chorizanthe represent a genus distinct from the annuals treated here (33 of the 41 annuals occur in the flora area, the rest are in Mexico or in South America). In that case, our annual members (if not submerged into Eriogonum) would be called Acanthogonum Torrey. The segregate genera allied to Chorizanthe, like those allied to Eriogonum, differ primarily in their involucres. Aristocapsa and Dodecahema, with haploid chromosome numbers of 14 and 17, respectively, are difficult to associate with any extant member of Chorizanthe (mainly n = 19, 20, 38, 40). Centrostegia, Lastarriaea, Mucronea, and Systenotheca all appear much more akin to Chorizanthe. A point of origin can be suggested only for Lastarriaea, namely Chorizanthe interposita Goodman of central Baja California, Mexico. C. D. Hardham (1989) reported a range of gametic numbers from single individuals. Until somatic counts are made, the primary chromosome numbers of some species remain uncertain. J. L. Reveal and C. B. Hardham (1989b) divided the annual species of Chorizanthe genus into three subgenera: subg. Amphietes (39 species), and subg. Eriogonella and subg. Quintaria (one species each). The first is divided into four sections (Ptelosepala, Acanthogonum, Fragile, and Clastoscapa Reveal & Hardham), all but the last of which are found in our region. Some sections were divided into subsections, which are not treated here. The approximately nine perennial species of subg. Chorizanthe occur only in arid regions of Chile and Argentina. (Discussion copyrighted by Flora of North America; reprinted with permission.) |
Varieties 2 (2 in the flora). (Discussion copyrighted by Flora of North America; reprinted with permission.) |
1. Involucres 3-5-toothed or 3-5-ribbed | → 2 |
1. Involucres 6-toothed (teeth sometimes vestigial or 6-ribbed) | → 7 |
2. Involucral teeth (4-)5 | → 3 |
| → 5 |
3. Involucral awns straight; perianths glabrous; filaments adnate at base of tepals; deserts, s California | C. spinosa |
3. Involucral awns uncinate; perianths pubescent abaxially; filaments adnate near top of floral tube; mountains, coastal mesas, deserts, w North America | → 4 |
4. Involucres campanulate, 1.5-2.5 mm, anterior tooth not leaflike; perianths white to rose, densely pubescent abaxially, 1.5-1.8(-2) mm; mountains and coastal mesas of California | C. polygonoides |
4. Involucres cylindric, 3-4.5 mm, anterior tooth leaflike; perianths yellow, thinly pubescent abaxially, 1.5-2.5 mm; deserts of w North America | C. watsonii |
5. Involucres cylindric, markedly transversely corrugate; perianths thinly pubescent abaxially; stamens 6 | C. corrugata |
5. Involucres urceolate to campanulate; perianths densely pubescent abaxially; stamens 9 | → 6 |
6. Plants erect; involucres urceolate, anterior tooth 5-10 mm with straight awn; achenes 3-gonous; warm deserts of sw North America | C. rigida |
6. Plants prostrate; involucres campanulate, anterior tooth 1.8-2 mm with uncinate awn; achenes lenticular; coastal mesas of extreme sw California | C. orcuttiana |
7. Involucral teeth with scarious or membranous margins | → 8 |
7. Involucral teeth without scarious or membranous margins | → 19 |
8. Involucral teeth margins membranous, continuous across sinuses | → 9 |
8. Involucral teeth margins scarious, parted or divided at sinuses, sometimes with distinct margin between erect teeth | → 12 |
9. Tepals obcordate to 2-lobed apically | → 10 |
9. Tepals entire apically | → 11 |
10. Tepals not denticulate apically; n, c California | C. stellulata |
10. Tepals denticulate apically; wc California | C. douglasii |
11. Involucres tomentose to floccose or glabrate, 3-4 mm; perianths densely pubescent abaxially, (1.5-)2.5-3 mm; sc Oregon and California | C. membranacea |
11. Involucres sparsely pubescent and hispid at least along ridges, 3-5 mm; perianths slightly pubescent abaxially, 3.5-4(-4.5) mm; wc California | C. douglasii |
12. Involucral awns straight | → 13 |
12. Involucral awns uncinate | → 15 |
13. Involucres congested in small bracteated clusters; leaf blades hirsute; tepals monomorphic; Coast Ranges of California | C. stellulata |
13. Involucres 1; leaf blades villous; tepals dimorphic; coastal dunes and grasslands of California | → 14 |
14. Perianths (3-)3.5-4.5 mm, pubescent nearly throughout; plants spreading or decumbent to somewhat erect | C. howellii |
14. Perianths (4-)5-6 mm, hairy on proximal 2; plants erect to spreading | C. valida |
15. Perianths glabrous; tepals entire apically; floral tubes lemon-yellow; coastal California | C. diffusa |
15. Perianths pubescent abaxially; floral tubes white | → 16 |
16. Involucres 2.5-4 mm, thinly pubescent abaxially; perianths 2.5-4 mm; wc California | C. robusta |
16. Involucres 1.5-2.5(-3) mm, villous abaxially; perianths 2-3 mm; coastal California | → 17 |
17. Tepals cuspidate apically | C. cuspidata |
17. Tepals erose apically | → 18 |
18. Involucral margins white to pink or purple; involucres 2-2.5(-3) mm; perianths 2-3.5 mm; stamens 9 | C. pungens |
18. Involucral margins pinkish; involucres 1.5-2(-2.5) mm; perianths 2-3 mm; stamens 3 or 6-9 | C. angustifolia |
19. Involucral awns unequal, anterior one greatly elongated | → 20 |
19. Involucral teeth of equal or alternating lengths but anterior tooth not greatly elongated | → 22 |
20. Anterior awns uncinate; c and n California | C. clevelandii |
20. Anterior awns straight; c California | → 21 |
21. Outer tepals obovate to nearly orbiculate, 2-lobed apically; inner tepals erose apically; stamens 9; Coast Ranges, c California | C. rectispina |
21. Outer tepals narrowly oblong, with minute cusp or 3 teeth apically; inner tepals entire apically; stamens 3; Coast Ranges, Transverse Ranges, Tehachapi Ranges, s Sierra Nevada, c California | C. uniaristata |
22. Tepals all, or at least inner ones, fimbriate or 2-lobed apically | → 23 |
22. Tepals entire, cuspidate, or erose apically | → 28 |
23. Outer and inner tepals fimbriate to laciniate apically; montane s California | C. fimbriata |
23. Outer tepals entire or if fimbriate to variously divided then inner tepals not fimbriate to laciniate; coastal ranges of California | → 24 |
24. Tepals white to pinkish | → 25 |
24. Tepals red, maroon, or dark purple | → 26 |
25. Inner tepals fimbriate apically | C. obovata |
25. Inner tepals 2-lobed apically | C. blakleyi |
26. Outer tepals entire, erect; involucres 3.5-4 mm; involucral awns 0.5-1 mm | C. palmeri |
26. Outer tepals shallowly lobed, obcordate, 2-lobed, erose, or at least wavy apically, spreading to recurved; involucres 4-6 mm; involucral awns 0.5-2 mm | → 27 |
27. Outer tepals 2-lobed or obcordate apically; inner tepals fimbriate apically; perianths (4.5-)5-6 mm; involucres 4-6 mm, slightly ventricose basally | C. biloba |
27. Outer tepals erose or at least wavy apically; inner tepals fimbriate or somewhat 2-lobed apically; perianths 4-4.5 mm; involucres 4-4.5 mm, strongly ventricose basally | C. ventricosa |
| → 29 |
28. Tepals dimorphic, sometimes monomorphic, inner ones usually narrower and shorter than outer ones | → 33 |
29. Plants spreading to erect; stems disarticulating at nodes; stamens 3; filaments adnate at top of floral tube | C. brevicornu |
29. Plants prostrate or decumbent to ascending or erect; stems not disarticulating at nodes; stamens 3 or 6-9 and filaments adnate at base of floral tube or stamens 6 or 9 and filaments adnate at top of floral tube | → 30 |
30. Filaments adnate at top of floral tube; tepals obtuse to truncate or minutely emarginate apically; achenes 3-gonous; mountains and coastal mesas of California | C. polygonoides |
30. Filaments adnate at base of floral tube; tepals entire, erose, or cuspidate apically; achenes lenticular or globose-lenticular; sw and wc California | → 31 |
31. Tepals entire apically; filaments connate basally; sw California | C. procumbens |
31. Tepals erose or cuspidate apically; filaments distinct; coastal c California | → 32 |
32. Tepals erose apically; flowers slightly exserted | C. angustifolia |
32. Tepals cuspidate apically; flowers included to slightly exserted | C. cuspidata |
33. Involucral awns straight; tepals erose to denticulate | → 34 |
33. Involucral awns uncinate; tepals entire or denticulate | → 35 |
34. Plants erect to spreading; involucres 3-4(-4.5) mm; wc California | C. valida |
34. Plants prostrate to spreading; involucres 1.5-2 mm; sw California | C. parryi |
35. Involucres 1.5-2 mm; s California | C. parryi |
35. Involucres 2-6 mm; California | → 36 |
36. Proximal leaflike bracts soon deciduous or absent | → 37 |
36. Proximal leaflike bracts persistent | → 38 |
37. Perianths 3-4(-5) mm; flowers mostly included; involucres usually congested terminally; coastal to montane sw California | C. staticoides |
37. Perianths 4.5-6 mm; flowers long-exserted; involucres 1; montane s California | C. leptotheca |
38. Perianths 4.5-6 mm; flowers long-exserted; inland California | C. xanti |
38. Perianths 2.5-3.5 mm; flowers exserted; coastal and insular sw California | → 39 |
39. Perianths 3-3.5 mm; involucres 2.5-3 mm, hairs slender, curly; stamens 9; coastal sw California | C. breweri |
39. Perianths 2.5-3 mm; involucres 2-2.5 mm, hairs stoutish, recurved; stamens 6; insular sw California | C. wheeleri |
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FNA vol. 5, p. 445. Author: James L. Reveal. |
FNA vol. 5, p. 461. |
Polygonaceae > subfam. Eriogonoideae |
Polygonaceae > subfam. Eriogonoideae > Chorizanthe > subg. Amphietes > sect. Ptelosepala |
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C. angustifolia, C. biloba, C. blakleyi, C. brevicornu, C. breweri, C. clevelandii, C. corrugata, C. cuspidata, C. diffusa, C. douglasii, C. fimbriata, C. howellii, C. leptotheca, C. membranacea, C. obovata, C. orcuttiana, C. palmeri, C. polygonoides, C. procumbens, C. pungens, C. rectispina, C. rigida, C. robusta, C. spinosa, C. staticoides, C. stellulata, C. uniaristata, C. valida, C. ventricosa, C. watsonii, C. wheeleri, C. xanti |
C. angustifolia, C. biloba, C. blakleyi, C. brevicornu, C. breweri, C. clevelandii, C. corrugata, C. cuspidata, C. diffusa, C. douglasii, C. fimbriata, C. howellii, C. leptotheca, C. membranacea, C. obovata, C. orcuttiana, C. palmeri, C. parryi, C. polygonoides, C. procumbens, C. pungens, C. rectispina, C. rigida, C. robusta, C. spinosa, C. staticoides, C. stellulata, C. uniaristata, C. valida, C. ventricosa, C. watsonii, C. wheeleri, C. xanti |
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R. Brown ex Bentham: Trans. Linn. Soc. London 17: 416, plate 17, fig. 11: plate 19. (1836) |
S. Watson: Proc. Amer. Acad. Arts 12: 271. (1877) |
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