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Bentham's monkeyflower, small-leaf monkey-flower

common monkey-flower, seep monkey-flower, yellow monkey-flower

Habit Annuals, fibrous-rooted. Perennials, rhizomatous, sometimes 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 to ascending-erect, branched distally, sometimes fistulose, to 10 mm wide, pressed, (6–)15–65(–80) cm, villous-glandular or moderately to densely hirtellous, hairs eglandular or glandular and eglandular.

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.

basal and cauline or basal not persistent;

petiole 0 mm or proximals 1–95 mm;

blade subpinnately, sometimes palmately, 5–7-veined, ovate-elliptic to ovate or suborbicular, 4–125 mm, 1–2 times longer than wide, gradually or abruptly reduced in size distally, base rounded to cuneate to truncate, margins crenate to coarsely dentate, proximally shallowly toothed to irregularly small-lobed or lyrate-dissected, apex rounded to obtuse, surfaces glabrous.

Flowers

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

herkogamous, (1–)3–20(–28), from distal nodes, sometimes in relatively compact racemes with reduced bracts.

Styles

sparsely hirtellous.

minutely 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, red-dotted, bilaterally symmetric, bilabiate;

tube-throat funnelform, (10–)12–20 mm, exserted 3–5 mm beyond calyx margin;

limb expanded 12–24 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.

15–40(–60) mm, villous-glandular or moderately to densely hirtellous, hairs eglandular or glandular and eglandular.

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.

nodding, usually without red markings, ovate-campanulate, inflated, sagittally compressed, 11–17(–20) mm, villous-glandular or moderately to densely hirtellous, hairs eglandular or glandular and eglandular, throat closing.

Capsules

included, 6–9(–11) mm.

included, 7–11(–12) mm.

Anthers

included, glabrous.

included, glabrous.

2n

= 28, 56.

= 28, 56.

Erythranthe microphylla

Erythranthe guttata

Phenology Flowering Mar–Jul. Flowering Apr–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. Springs and seeps, marshes, beaver dams, along rivers, streams, and irrigation canals, loamy soils in conifer forests, wet and damp meadows, wet roadsides.
Elevation 20–1700(–2600) m. (100–5600(–8500) ft.) 20–3200(–3700) m. (100–10500(–12100) ft.)
Distribution
from FNA
CA; ID; NV; OR; WA; BC
[WildflowerSearch map]
from FNA
AK; AZ; CA; CO; CT; ID; MI; MT; NE; NM; NV; NY; OR; PA; SD; TX; UT; WA; WY; AB; BC; NB; NT; SK; YT; Mexico (Baja California, Baja California Sur, Chihuahua, Coahuila, Nayarit, Sonora) [Introduced in Europe]
[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.)

Erythranthe guttata is markedly variable in stature, leaf shape, vestiture, flower size, and the separation distance between anthers and stigma; it ranges from subalpine and near-alpine habitats into desert situations where water is available.

In all of Colorado, the Four Corners area, and north-central New Mexico, the vestiture of stems and calyces is consistently densely hirsute-hirtellous, without glandular hairs. Plants with similar vestiture also occur in British Columbia, Oregon, and Washington, and in scattered localities elsewhere. In northwestern Arizona, California Nevada, and southern Oregon, vestiture is consistently villous-glandular, without eglandular hairs. Elsewhere in the geographic range the vestiture is a mix of hirsute-hirtellous (eglandular) and villous-glandular hairs. Other morphological variants and patterns, as well as variation in ploidy level, within Erythranthe guttata were discussed by G. L. Nesom (2012i).

Plants of Erythranthe guttata with extremely large corollas have been frequently collected on the Aleutian Islands, Kodiak Island, and in other Alaskan localities (for example, Admiralty Island Amakuk, Juneau, and Yakutat Bay). Corolla tube-throats are 19–26 mm, and the limbs are expanded to 18–25 mm. The type collection of E. guttata is one of these plants, and the name E. guttata may prove to apply most appropriately only to Alaskan populations. Diploids and tetraploids appear to be sympatric in Alaska.

Mimulus guttatus subsp. haidensis was described as an endemic subalpine race that occurs in and along the flanks of the Queen Charlotte Mountains on Graham Island and Moresby Island. The subspecies was distinguished on the basis of its hirtellous vestiture, but plants of similar hirtellous vestiture occur over the whole range of the species. A tetraploid chromosome number (2n = 56) was reported for subsp. haidensis from a total of five localities on Graham Island and Moresby Island (J. A. Calder and R. L. Taylor 1968, vol. 2), and diploids (2n = 28) were documented from one locality on each of the two islands. At least one of the diploids has densely hirtellous stems, pedicels, and calyces, matching the morphology of subsp. haidensis.

Erythranthe guttata is naturalized in Europe and has been introduced to the northeastern United States (Connecticut, Michigan, New York, and Pennsylvania) and eastern Canada (New Brunswick).

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

Source FNA vol. 17, p. 415. FNA vol. 17, p. 411.
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. 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
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 guttatus, M. clementinus, M. equinus, M. glabratus var. adscendens, M. grandiflorus, M. guttatus subsp. haidensis, M. guttatus var. puberulus, M. hirsutus, M. langsdorffii var. argutus, M. langsdorffii var. californicus, M. langsdorffii var. guttatus, M. langsdorffii var. minimus, M. langsdorffii var. platyphyllus, M. lyratus, M. paniculatus, M. petiolaris, M. prionophyllus, M. puberulus, M. rivularis
Name authority (Bentham) G. L. Nesom: Phytoneuron 2012-39: 44. (2012) (de Candolle) G. L. Nesom: Phytoneuron 2012-39: 43. (2012)
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