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bent-stem monkeyflower, Dudley's monkeyflower

Bentham's monkeyflower, small-leaf monkey-flower

Habit Annuals, fibrous-rooted or filiform-taprooted. Annuals, fibrous-rooted.
Stems

ascending to decumbent or prostrate, geniculate at nodes, simple or diffusely branched, 5–60 cm, moderately villous, hairs 0.8–2 mm, multicellular, eglandular and also 0.1–0.3 mm, stipitate-glandular.

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.

Leaves

basal and cauline, basal usually deciduous by flowering;

petiole 2–10(–35) mm;

blade pinnately to subpinnately veined, broadly ovate or elliptic-ovate to triangular, 8–35 × 5–30 mm, base cuneate to rounded or subcordate, margins serrate or dentate, teeth 3–10 per side, apex acute to obtuse or rounded, surfaces moderately villous, hairs 0.8–2 mm, multicellular, eglandular, and 0.1–0.3 mm, stipitate-glandular.

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.

Flowers

herkogamous, (1–)6–20, from all or medial to distal nodes.

herkogamous, 1–8(–14), usually from distal nodes, chasmogamous.

Styles

glabrous.

sparsely hirtellous.

Corollas

yellow, without white patches, throat red-spotted, spots concentrated or becoming coalescent into a somewhat discrete splotch at base of each of 3 abaxial lobes and sometimes 2 adaxial, bilaterally symmetric, ± bilabiate;

tube-throat cylindric, 9–12 mm, exserted beyond calyx margin;

limb expanded 10–18 mm diam.

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.

Fruiting pedicels

12–26(–55) mm, moderately villous, hairs 0.8–2 mm, multicellular, eglandular and also 0.1–0.3 mm, stipitate-glandular.

8–30(–50) mm, distals hirtellous, hairs commonly deflexed, or mixed hirtellous and stipitate-glandular, sometimes only short villous-glandular, hairs gland-tipped.

Fruiting calyces

red-spotted, campanulate-cylindric, weakly inflated, (5–)6–8 mm, margins distinctly toothed or lobed, sparsely to moderately villous-glandular, ribs shallowly wing-angled, lobes pronounced, erect to spreading or spreading-recurving.

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.

Capsules

included, 4–6(–7) mm.

included, 6–9(–11) mm.

Anthers

included, glabrous.

included, glabrous.

2n

= 32.

= 28, 56.

Erythranthe geniculata

Erythranthe microphylla

Phenology Flowering (Mar–)Apr–Jul. Flowering Mar–Jul.
Habitat Granite crevices, canyon slopes, talus, crevices in volcanic outcrops, edges of boulders, roadsides, damp sandy soils, sandy water edges, gravelly soils and creek bottoms. 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.
Elevation 200–900(–1200) m. (700–3000(–3900) ft.) 20–1700(–2600) m. (100–5600(–8500) ft.)
Distribution
from FNA
CA
[WildflowerSearch map]
from FNA
CA; ID; NV; OR; WA; BC
[WildflowerSearch map]
Discussion

Erythranthe geniculata is known from an apparently disjunct cluster of populations in Butte, Sutter, and Yuba counties and then from Tuolumne and Stanislaus counties south to Kern County.

Erythranthe geniculata, compared to E. floribunda, has larger, chasmogamous, and allogamous flowers. The anther pairs of E. geniculata are at different levels, and the stigma is slightly above the adaxial anther pair; in E. floribunda both anther pairs and the stigma are at the same level.

Erythranthe arenaria, E. geniculata, and E. norrisii constitute a group of apparently closely related species within sect. Mimulosma endemic along the Sierra Nevada. All have ovate-petiolate leaves (only the basal ones are sometimes ovate in E. arenaria) with pinnate to subpinnate venation. The more widespread E. floribunda, which is part of the above group, also is similar, but all three endemics have larger corollas with the tube exserted at greater length beyond the calyx margin.

(Discussion copyrighted by Flora of North America; reprinted with permission.)

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.)

Source FNA vol. 17, p. 404. FNA vol. 17, p. 415.
Parent taxa Phrymaceae > Erythranthe Phrymaceae > Erythranthe
Sibling taxa
E. acutidens, E. alsinoides, E. ampliata, E. androsacea, E. arenaria, E. arenicola, E. arvensis, E. barbata, E. bicolor, E. brachystylis, E. breviflora, E. breweri, E. caespitosa, E. calcicola, E. calciphila, E. cardinalis, E. carsonensis, E. charlestonensis, E. chinatiensis, E. cinnabarina, E. corallina, E. cordata, E. decora, E. dentata, E. diffusa, E. discolor, E. eastwoodiae, E. erubescens, E. exigua, E. filicaulis, E. filicifolia, E. floribunda, E. gemmipara, E. geyeri, E. glaucescens, E. gracilipes, E. grandis, E. grayi, E. guttata, E. hallii, E. hardhamiae, E. hymenophylla, E. inamoena, E. inconspicua, E. inflatula, E. jungermannioides, E. laciniata, E. latidens, E. lewisii, E. linearifolia, E. marmorata, E. michiganensis, E. microphylla, E. minor, E. montioides, E. moschata, E. nasuta, E. norrisii, E. nudata, E. palmeri, E. pardalis, E. parishii, E. parvula, E. patula, E. percaulis, E. primuloides, E. ptilota, E. pulsiferae, E. purpurea, E. regni, E. rhodopetra, E. rubella, E. scouleri, E. shevockii, E. sierrae, E. suksdorfii, E. taylorii, E. thermalis, E. tilingii, E. trinitiensis, E. unimaculata, E. utahensis, E. verbenacea, E. washingtonensis, E. willisii
E. acutidens, E. alsinoides, E. ampliata, E. androsacea, E. arenaria, E. arenicola, E. arvensis, E. barbata, E. bicolor, E. brachystylis, E. breviflora, E. breweri, E. caespitosa, E. calcicola, E. calciphila, E. cardinalis, E. carsonensis, E. charlestonensis, E. chinatiensis, E. cinnabarina, E. corallina, E. cordata, E. decora, E. dentata, E. diffusa, E. discolor, E. eastwoodiae, E. erubescens, E. exigua, E. filicaulis, E. filicifolia, E. floribunda, E. gemmipara, E. geniculata, E. geyeri, E. glaucescens, E. gracilipes, E. grandis, E. grayi, E. guttata, E. hallii, E. hardhamiae, E. hymenophylla, E. inamoena, E. inconspicua, E. inflatula, E. jungermannioides, E. laciniata, E. latidens, E. lewisii, E. linearifolia, E. marmorata, E. michiganensis, E. minor, E. montioides, E. moschata, E. nasuta, E. norrisii, E. nudata, E. palmeri, E. pardalis, E. parishii, E. parvula, E. patula, E. percaulis, E. primuloides, E. ptilota, E. pulsiferae, E. purpurea, E. regni, E. rhodopetra, E. rubella, E. scouleri, E. shevockii, E. sierrae, E. suksdorfii, E. taylorii, E. thermalis, E. tilingii, E. trinitiensis, E. unimaculata, E. utahensis, E. verbenacea, E. washingtonensis, E. willisii
Synonyms Mimulus geniculatus, M. dudleyi, M. floribundus var. geniculatus 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
Name authority (Greene) G. L. Nesom: Phytoneuron 2012-39: 38. (2012) (Bentham) G. L. Nesom: Phytoneuron 2012-39: 44. (2012)
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