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Bentham's monkeyflower, small-leaf monkey-flower

Columbia River monkey-flower, Scouler's monkey-flower

Habit Annuals, fibrous-rooted. Perennials, rhizomatous, producing long, sparsely leafy runners from basal nodes.
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, simple or few-branched, 15–80 cm, glabrous.

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, basal usually not persistent;

petiole: proximals and medial 10–25 mm (gradually merging into blade), distals 0 mm;

blade palmately (3–)5–7-veined to subpinnate, (proximal to medial) oblong-elliptic to oblong-lanceolate, 25–60 × 8–18 mm, usually 3–4 times longer than wide, base attenuate, margins evenly, shallowly dentate or crenate to mucronate or mucronulate, teeth 10–20 per side, sometimes more deeply toothed at base, apex obtuse to acute, surfaces glabrous.

Flowers

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

herkogamous, (1 or)2–8, from distal nodes.

Styles

sparsely hirtellous.

minutely, prominently hirsutulous to villosulous.

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, without red markings, bilaterally symmetric, bilabiate;

tube-throat funnelform, 20–24 mm, exserted 10–15 mm beyond calyx margin;

limb expanded 22–30 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.

20–25 mm, glabrous.

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.

ovoid, inflated, sagittally compressed, 13–14 mm, glabrous, throat closing.

Capsules

included, 6–9(–11) mm.

unknown.

Anthers

included, glabrous.

included, glabrous.

2n

= 28, 56.

Erythranthe microphylla

Erythranthe scouleri

Phenology Flowering Mar–Jul. Flowering May–Aug.
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. Salt marshes, river banks.
Elevation 20–1700(–2600) m. (100–5600(–8500) ft.) 0–20 m. (0–100 ft.)
Distribution
from FNA
CA; ID; NV; OR; WA; BC
[WildflowerSearch map]
from FNA
OR; WA
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.)

Erythranthe scouleri is distinctive in its oblong-elliptic leaves with long-tapering bases and closely toothed margins, completely glabrous vestiture, long, leafy runners from basal cauline nodes, large corollas with broad limbs, and prominently hairy styles. Several features suggest a close relationship to E. decora, particularly its very large corollas, hairy styles, closely toothed leaf margins, tall, simple, and erect stems, numerous runners, and its geographic range. All collections apparently have been made near the mouth of the Columbia River in Clatsop and Columbia counties, Oregon (G. L. Nesom 2013d). Recent observations (Alexander John Wright, pers. comm.; photos http://www.inaturalist.org/observations/3537008) indicate that it also occurs in Wahkiakum County, Washington, in the Julia Butler Hansen Refuge.

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

Source FNA vol. 17, p. 415. 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. 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
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. 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. 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 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 scouleri, M. guttatus subsp. scouleri
Name authority (Bentham) G. L. Nesom: Phytoneuron 2012-39: 44. (2012) (Hooker) G. L. Nesom: Phytoneuron 2012-39: 44. (2012)
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