Erythranthe microphylla |
Erythranthe tilingii |
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Bentham's monkeyflower, small-leaf monkey-flower |
larger mountain monkeyflower, mountain monkey-flower, Tiling's monkey-flower |
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Habit | Annuals, fibrous-rooted. | Perennials, rhizomatous, solitary to weakly colonial, rhizomes forming a mass, yellowish, branching, filiform. |
Stems | erect, simple, sometimes many-branched from basal cauline nodes, terete, sometimes distinctly 4-angled, (3–)5–30(–45) cm, glabrous below inflorescence, sometimes distals hirtellous, hairs commonly deflexed, or mixed hirtellous and stipitate-glandular, sometimes only short villous-glandular, hairs gland-tipped. |
erect-ascending, usually freely branched, 2–35 cm, glabrous or sparsely stipitate-glandular to short glandular-villous. |
Leaves | basal and cauline, basal sometimes absent at flowering; petiole: basal or proximals to medials 3–25(–35) mm, distals 0 mm (then blade subclasping to narrowly perfoliate); blade often purplish, palmately 3–5-veined, ovate or ovate-lanceolate to elliptic-ovate, suborbicular, or depressed-ovate, (3–)10–35 × 3–25 mm, base rounded to truncate or subcordate, margins shallowly crenate to sharply crenate-serrate, teeth 5–10 per side, basal and proximal often irregularly incised near petiole and sublyrate, apex acute to obtuse-rounded, surfaces glabrous or sparsely to moderately hirtellous, eglandular. |
cauline; petiole 0–25 mm, distals 0 mm; blade palmately 3–5-veined, ovate to lanceolate-triangular or narrowly lanceolate (broadly ovate in large-leaved forms), 5–35(–55) mm, base cuneate to attenuate, margins irregularly denticulate, apex acute to obtuse or rounded, surfaces glabrous, sparsely stipitate-glandular to short glandular-villous, glabrate, or sparsely to moderately villous, hairs thick-vitreous, eglandular. |
Flowers | herkogamous, 1–8(–14), usually from distal nodes, chasmogamous. |
herkogamous, 1–3(–5), from distal nodes. |
Styles | sparsely hirtellous. |
hirtellous. |
Corollas | yellow to golden yellow or orangish yellow, usually red-spotted, abaxial limb sometimes with a large red splotch, bilaterally symmetric, bilabiate; tube-throat broadly funnelform, (6–)8–16(–20) mm, exserted (1–)2–6(–8) mm beyond calyx margin; limb expanded 8–25 mm, palate villous. |
yellow, red-dotted, bilaterally symmetric, bilabiate; tube-throat broadly funnelform, 15–28 mm, exserted 5–10 mm beyond calyx margin; limb expanded 14–30 mm. |
Fruiting pedicels | 8–30(–50) mm, distals hirtellous, hairs commonly deflexed, or mixed hirtellous and stipitate-glandular, sometimes only short villous-glandular, hairs gland-tipped. |
15–35(–40) mm, sparsely stipitate-glandular to short glandular-villous. |
Fruiting calyces | nodding 30–90º, sometimes red-tinged or -dotted, ovoid-campanulate to broadly cylindric-campanulate, inflated, sagittally compressed, (7–)9–16(–20) mm, minutely hirtellous, hairs sometimes reduced to basal cells, or glabrous, throat strongly to weakly closing. |
usually purple-tinged and purple-dotted, broadly campanulate, inflated, sagittally compressed, 11–15 mm, glabrous or sparsely stipitate-glandular to short glandular-villous, villous at sinuses, throat closing, lobes broadly ovate, abaxial usually longer than lateral, adaxial at least 2 times as long as others. |
Capsules | included, 6–9(–11) mm. |
included, 5–7 mm. |
Anthers | included, glabrous. |
included, glabrous. |
2n | = 28, 56. |
= 28, 56. |
Erythranthe microphylla |
Erythranthe tilingii |
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Phenology | Flowering Mar–Jul. | Flowering Jul–Sep. |
Habitat | Rock depressions, rocky ridges, cliff faces, roadcuts, wet meadows, seeps, stream banks, drying ponds, ephemeral stream channels, vernal springs over serpentine, roadsides and roadside ditches, dry banks, lava soils, loam, clay, gravel, yellow pine, oak-pine, mixed oak woodlands, oak-chaparral. | Seeps, springs, stream banks, shallow rivulets, cliff bases, ledges and crevices, steep gravelly slopes, wet meadows. |
Elevation | 20–1700(–2600) m. (100–5600(–8500) ft.) | 1400–3400 m. (4600–11200 ft.) |
Distribution |
CA; ID; NV; OR; WA; BC
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AZ; CA; ID; MT; NV; OR; UT; AB
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Discussion | Erythranthe microphylla is characterized by its annual duration (fibrous-rooted), usually simple stems, relatively widely spaced leaves, glabrous or hirtellous vestiture, open corollas, and calyces closing at the throat. Even in the smallest corollas, the stigma is positioned above the adaxial anther pair, indicating that all are primarily allogamous. Some plants have basal and proximal cauline leaves with exaggeratedly and irregularly toothed-incised margins (especially in Lake and Napa counties, California, as in the types of Mimulus glareosus and M. guttatus var. insignis, respectively), but a similar tendency can be seen over most of the geographic range. Plants of E. microphylla vary greatly in height, leaf size, and flower size (the larger flowers approaching the size of those in E. grandis and E. decora), yet all seem to be within the expression of a single species. An inversion sequence of chromosome 8 (the DIV1 region) is perfectly correlated with the life history features of at least four species of sect. Simiolus (D. B. Lowry and J. H. Willis 2010). One sequence occurs in Erythranthe guttata and E. grandis, which are perennial and rhizomatous, occur in habitats with year-round moisture, and flower relatively late in the season, while the opposite sequence occurs in E. nasuta and E. microphylla, which are annual and slender-taprooted or fibrous-rooted, occur in quickly drying habitats, and flower in early season. The inversion, with its tightly linked, locally adaptive alleles, contributes to isolating mechanisms between species with contrasting sequences and preserves the constellation of features that makes each a recognizable entity. Hybridization and apparent introgression occur among Erythranthe guttata, E. microphylla, and E. nasuta, yet each remains distinct in duration, perennating morphology, and habit. Erythranthe arvensis (annual) also is a member of this gene-sharing group. A whole-genome analysis by A. D. Twyford and J. Friedman (2015) showed that populations of E. guttata and E. microphylla cluster together within each of several geographically delimited regions; they interpreted their trees as showing phylogenetic relationships and concluded that E. guttata and E. microphylla are conspecific. Other evidence (G. L. Nesom 2014c, 2014d), particularly the DIV1 inversion sequence, indicates that E. guttata and E. grandis are most closely related to each other and were derived from an ancestor of annual duration. Thus, E. microphylla and E. nasuta are morphologically and phylogenetically distinct from E. guttata, despite extensive gene flow where they are sympatric with it. (Discussion copyrighted by Flora of North America; reprinted with permission.) |
Plants of Erythranthe tilingii are characterized by their relatively low stature and stems arising from a system of thin rhizomes and producing mostly one to three large flowers each; they usually occur at relatively high elevations. Erythranthe tilingii sometimes has been considered to include one or several infraspecific entities; from within this taxonomic amalgam, four distinct species are recognized here: E. caespitosa, E. corallina, E. minor, and E. tilingii. Erythranthe corallina and E. minor probably are more closely related to E. guttata. The populations identified here as E. tilingii from northeastern Oregon northeast to Alberta and southeast to Utah may prove to be a separate (undescribed) species. Erythranthe tilingii in the strict sense is relatively widespread over the western United States and is sympatric with E. caespitosa and E. corallina. Leaves in E. tilingii are variable in size, and particularly in Idaho, they may approach the small size of those of E. caespitosa, but the leaf margins of E. tilingii are distinctly toothed, and the stems are taller and more erect. Across the range of the species, plants sometimes produce very large leaves, but these often occur on plants with characteristically smaller leaves. This wide variability in size apparently does not occur in E. caespitosa. (Discussion copyrighted by Flora of North America; reprinted with permission.) |
Source | FNA vol. 17, p. 415. | FNA vol. 17, p. 408. |
Parent taxa | Phrymaceae > Erythranthe | Phrymaceae > Erythranthe |
Sibling taxa | ||
Synonyms | Mimulus microphyllus, M. glareosus, M. guttatus var. depauperatus, M. guttatus var. insignis, M. guttatus var. microphyllus, M. guttatus var. platycalyx, M. langsdorffii var. insignis, M. langsdorffii var. microphyllus, M. luteus var. depauperatus, M. nasutus var. insignis, M. platycalyx | Mimulus tilingii, M. caespitosus var. implexus, M. implexus, M. implicatus, M. langsdorffii var. tilingii, M. lucens, M. veronicifolius |
Name authority | (Bentham) G. L. Nesom: Phytoneuron 2012-39: 44. (2012) | (Regel) G. L. Nesom: Phytoneuron 2012-39: 44. (2012) |
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