Keckiella |
Keckiella ternata |
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bush penstemon, keckiella, penstemon |
blue stem keckiella, scarlet keckiella |
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Habit | Subshrubs or shrubs. | |||||||||||||||||||||||||||||
Stems | erect to spreading, sometimes climbing, glabrous or hairy. |
spreading to erect, 5–25 dm, glabrous when young, glaucous. |
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Leaves | drought-deciduous, cauline, opposite, subopposite, or whorled; petiole present or absent; blade fleshy or not, leathery or not, margins entire or toothed, sometimes relatively small, distal leaf blade not needlelike or scalelike. |
whorled (3s), sometimes opposite; blade linear to narrowly oblanceolate or lanceolate, 15–60 mm, margins 4–10-toothed. |
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Inflorescences | terminal, panicles, thyrses, corymbs, or spikelike racemes; bracts present. |
panicles, glabrous or glandular-hairy. |
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Pedicels | present; bracteoles present, sometimes much reduced. |
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Flowers | bisexual; sepals 5, basally connate, calyx weakly bilaterally symmetric, short-tubular, lobes lanceolate to ovate, sometimes oblanceolate; corolla red, reddish orange, pink, white, cream, yellow, brownish yellow, or purplish brown, sometimes with red-purple nectar guides, bilaterally symmetric, strongly bilabiate, ± tubular, tube base not spurred or gibbous, lobes 5, abaxial 3, adaxial 2, adaxial lip hooded; stamens 4, basally adnate to corolla, didynamous, filaments basally eglandular-hairy, pollen sacs explanate; staminode 1, filamentous; nectary a hypogynous disc; ovary 2-locular, placentation axile; stigma capitate. |
calyx 3.8–7.2 mm, lobes lanceolate to ovate; corolla red, 21–31 mm, tube plus indistinct throat 16–24 mm, adaxial lip 5.5–9.5 mm; pollen sacs 0.8–1.1 mm; staminode densely yellow-hairy, included. |
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Fruits | capsules, dehiscence septicidal, sometimes also loculicidal distally, not densely packed with white membranous hairs. |
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Seeds | 20–100, brown, ovoid, wings essentially absent. |
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x | = 8. |
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Keckiella |
Keckiella ternata |
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Distribution |
w United States; nw Mexico |
CA; nw Mexico
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Discussion | Species 7 (7 in the flora). Keckiella is nearly endemic to the California Floristic Province (P. H. Raven and D. I. Axelrod 1978). In the flora area, taxa range beyond California into Arizona and Nevada; two occur in Mexico (Baja California Peninsula). All taxa of Keckiella were first described in Penstemon, where they were long recognized as related to each other and anomalous within Penstemon. Keckiella taxa were treated as Keckia Straw before the prior use of that name for a fossil alga was noticed. The relationship of Keckiella and Penstemon with the rest of Cheloneae was clarified by A. D. Wolfe et al. (2002). This confirmed the classification implied by D. D. Keck (1936) and proposed by R. M. Straw (1966). The two genera are now recognized as distinct evolutionary lineages based on morphology, rust fungi relationships, phytochemistry, and DNA evidence (D. B. O. Savile 1968b, 1979; D. C. Michener 1982; O. Mistretta and R. Scogin 1989; Wolfe et al.; C. E. Freeman et al. 2003). Among Keckiella species, gross morphological and anatomical characteristics of leaf and wood correlate with habitat; floral characters, other than the distinctive nectary structure that helps define Keckiella, reflect the pollination syndrome (D. C. Michener 1981, 1982). Leaf morphology is typical of drought-deciduous plants (Michener 1981). Varieties are recognized in part on foliage and calyx indument. Leaf primordia of all species are (weakly) pubescent with glandular and non-glandular hairs, making indument a challenging character for interpretation. Pollination biology and habitat tolerances appear to have been important in species diversification. Bee pollination is fundamental in Keckiella, with floral size among the species scaled to a range of native bees; hummingbird pollination is polyphyletic in parallel with Penstemon (D. C. Michener 1982; C. E. Freeman et al. 2003; Paul Wilson et al. 2007). Natural hybrids have been found repeatedly for two species pairs (K. antirrhinoides var. antirrhinoides and K. cordifolia; K. breviflora var. glabrisepala and K. lemmonii) where the ranges overlap. The pollination syndromes are not effective barriers to extraneous cross pollination: even the hummingbird-pollinated species are visited by small pollen-foraging bees that may promote some pollination or hybridization separate from the hummingbirds (Michener). Leaf flavonoids do not support the hummingbird-pollinated species being each other’s closest relatives (O. Mistretta and R. Scogin 1989). Species of Keckiella are widespread but only locally common in their habitat. They usually occur in unstable sites such as slopes, scree, and crevices in rocky faces from near sea level to 3000 m. These local micro-sites may be less fire prone than the immediately surrounding communities. Fire ecology may be important in the evolution of the species; root gnarls found only in K. antirrhinoides var. antirrhinoides and K. cordifolia may help these species resprout following the fires typical of their low-elevation habitats (D. C. Michener 1981). The eco-evolutionary segregation of Keckiella from its relatives may have been centered on the Klamath region during the Tertiary (A. D. Wolfe et al. 2002). C. E. Freeman et al. (2003) recognized three clades within Keckiella: the basal-most K. rothrockii lineage, a northern clade (K. breviflora, K. corymbosa, K. lemmonii, and K. ternata), ranging from southern to northern California and through almost the entire 3000 m elevational range of the genus, and a southern clade (K. antirrhinoides and K. cordifolia) nearly restricted to lower elevations in southern California and Baja California, Mexico. They argued that speciation in these clades likely reflects repeated regional and elevational migration during the Pleistocene, presumably including the ecological scenarios that led to the modern species in fire-dominated vegetation types. (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.) |
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Key |
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Source | FNA vol. 17, p. 75. | FNA vol. 17, p. 79. | ||||||||||||||||||||||||||||
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Synonyms | Lepidostemon | Penstemon ternatus | ||||||||||||||||||||||||||||
Name authority | Straw: Brittonia 19: 203. (1967) | (Torrey e× A. Gray) Straw: Brittonia 19: 204. (1967) | ||||||||||||||||||||||||||||
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