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Bentham's monkeyflower, small-leaf monkey-flower

mimule musqué, musk monkeyflower, musk-flower, musk-plant, sticky monkey-flower

Habit Annuals, fibrous-rooted. Perennials, rhizomatous, rooting at proximal 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, sometimes ascending to decumbent, simple or branched, (2–)5–20 cm, nodes 2–4(or 5), glabrate to glandular-villous, hairs 0.5–2 mm, gland-tipped, internodes evident.

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.

usually cauline, basal not persistent, distinctly separated;

petiole 0 mm or (0.5–)1–5(–10) mm;

blade pinnately veined, oblong-ovate to ovate, (10–)15–40(–50) × 5–25 mm, base obtuse-cuneate to truncate, rounded or subcordate, subclasping to sessile, margins coarsely serrate-dentate to denticulate or subentire, apex acute to obtuse, surfaces glabrate to glandular-villous.

Flowers

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

herkogamous, 1–8, from medial to distal nodes.

Styles

sparsely hirtellous.

glabrous.

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, throat with fine red to blackish or brown lines extending onto lobes, red to brown dots in throat and lobes present or absent, bilaterally or nearly radially symmetric, bilabiate or nearly regular;

tube-throat narrowly funnelform, 11–18 mm, exserted beyond calyx margin;

lobes oblong-obovate, apex rounded to notched.

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.

(7–)10–25 mm, glabrate to 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.

ridge- to wing-angled, campanulate to cylindric-campanulate, weakly or not inflated, 6–13 mm, villous to glandular-villous, lobes erect to spreading-recurving, strongly unequal to subequal, triangular to linear-lanceolate or narrowly triangular-acuminate, 2–4 mm, apex acute to obtuse.

Capsules

included, 6–9(–11) mm.

included, 6–8 mm.

Anthers

included, glabrous.

included, glabrous or slightly hirtellous to scabrous.

2n

= 28, 56.

= 32.

Erythranthe microphylla

Erythranthe moschata

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. Springs and seeps, creek edges, moist meadows, ditches, along trails, roadsides, rocky ridges, granite outcrops, shaded and wet places in sagebrush, aspen, fir, spruce-fir, lodgepole pine forests, meadows.
Elevation 20–1700(–2600) m. (100–5600(–8500) ft.) (300–)400–3100 m. ((1000–)1300–10200 ft.)
Distribution
from FNA
CA; ID; NV; OR; WA; BC
[WildflowerSearch map]
from FNA
CA; CO; CT; ID; MA; ME; MI; MT; NH; NJ; NV; NY; OR; PA; RI; UT; VA; VT; WA; WI; WV; WY; BC; NB; NF; NS; ON; PE; QC; SPM [Introduced in South America (Chile), Europe, e Asia (Japan), Pacific Islands (New Zealand), Australia]
[WildflowerSearch map]
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.)

Earlier segregation of Erythranthe moniliformis as distinct from E. moschata (for example, G. L. Nesom 2012g) emphasized a primarily erect habit and tendency toward sessile to subsessile and more densely arranged cauline leaves in E. moniliformis versus a decumbent to procumbent habit and consistently petiolate leaves on longer internodes in E. moschata. Discontinuities in morphology, geography, and ecology were not confirmed in later study by Nesom (2017). Rhizomes with small, tuberlike swellings can be observed over the whole moschata/moniliformis range, and there apparently are no consistent distinctions in vestiture and corolla size.

Mimulus acutidens Reiche (1911), a later homonym of M. acutidens Greene, pertains here.

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

Source FNA vol. 17, p. 415. FNA vol. 17, p. 401.
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. 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 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 moschatus, E. inodora, E. moniliformis, M. crinitus, M. guttatus var. moschatus, M. inodorus, M. leibergii, M. macranthus, M. moniliformis, M. moschatus var. longiflorus, M. moschatus var. moniliformis, M. moschatus var. pallidiflorus
Name authority (Bentham) G. L. Nesom: Phytoneuron 2012-39: 44. (2012) (Douglas ex Lindley) G. L. Nesom: Phytoneuron 2012-39: 38. (2012)
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