FNA: Erythranthe verbenacea vs. Erythranthe willisii

	Erythranthe verbenacea	Erythranthe willisii
	crimson monkeyflower, pico Pajaro	Willis' monkeyflower
Habit	Perennials, rhizomatous.	Perennials, rhizomatous, rarely rooting at proximal nodes, usually forming large colonies, rhizomes white, usually highly branching.
Stems	erect to decumbent, usually simple, weakly 4-angled, 20–60 cm, ± glandular-villous.	usually sprawling-decumbent, branched, sometimes simple, 7–45 cm, nodes (2–)4–15+, densely glandular-villous, hairs 1–2 mm, glandular, internodes evident.
Leaves	cauline; petiole 0 mm; blade palmately 3–5-veined, elliptic to obovate, rhombic-ovate, or broadly spatulate, 50–75 × 15–26(–30) mm, base subcordate, subclasping, margins coarsely serrate, sometimes only distally, apex acute to obtuse, surfaces ± glandular-villous.	usually cauline, basal not persistent, distinctly separated; petiole 0 mm, sometimes 1–2 mm at proximal nodes; blade bicolored, purplish abaxially, pinnately veined, ovate to elliptic-ovate, midcauline 10–35 × 6–18 mm, base rounded to subcordate, margins coarsely serrate-dentate to denticulate or subentire, apex short-attenuate to acute, obtuse, or rounded, surfaces densely glandular-villous, hairs 1–2 mm, gland-tipped.
Flowers	herkogamous, 2–12, axillary at leafy medial to distal nodes.	herkogamous, (4–)8–30+, from medial to distal nodes, sometimes from all nodes.
Styles	glabrous.	glabrous.
Corollas	crimson, often yellow-tinged, palate ridges dark red, bilaterally symmetric, strongly bilabiate; tube-throat tubular, 25–35 mm, exserted 13–25 mm beyond calyx margin; abaxial limb spreading, adaxial erect, lobe apex truncate, often emarginate, throat open, palate ridges densely short-villous.	yellow, throat, tube, and proximal portion of abaxial 3 lobes with fine, red to brownish lines, weakly bilaterally or nearly radially symmetric, weakly bilabiate or nearly regular; tube-throat narrowly funnelform, 12–15 mm, exserted beyond calyx margin; limb 9–12 mm wide (pressed), lobes oblong-obovate, apex rounded to notched.
Fruiting pedicels	45–90(–150) mm.	4–20(–25) mm, densely glandular-villous, hairs 1–2 mm, gland-tipped.
Fruiting calyces	campanulate, weakly inflated, 20–28 mm, sparsely glandular-villosulous to stipitate-glandular, lobes triangular to ovate-triangular, apex linear-triangular.	ridge- to wing-angled, campanulate to cylindric-campanulate, weakly inflated, 7–10 mm, densely glandular-villous, lobes erect to slightly spreading, unequal, triangular to linear-lanceolate, 2–4 mm, apex acuminate-apiculate.
Capsules	included, 15–22 mm.	included, 4–5 mm.
Anthers	exserted, white-villous, thecae reflexed 45º.	included, glabrous or finely hirtellous to scabrous.
2n	= 16.

	Erythranthe verbenacea	Erythranthe willisii
Phenology	Flowering Jun–Sep.	Flowering May–Sep.
Habitat	Stream edges and beds, flood plains, around seeps and springs, canyon bottoms, moist cliff crevices and ledges.	Seepage, drainage margins, moist soils, talus, cracks and crevices, soils deprived from serpentine.
Elevation	300–2600 m. (1000–8500 ft.)	(500–)700–900 m. ((1600–)2300–3000 ft.)
Distribution	AZ; UT; Mexico (Baja California, Chihuahua, Durango, Sinaloa, Sonora) [WildflowerSearch map]	CA
Discussion	R. K. Vickery (1992) noted that yellow-flowered morphs of Erythranthe verbenacea occur in a population at Vasey’s Paradise in the Grand Canyon (Coconino County), 32 miles downstream from Lees Ferry. Populations of Erythranthe verbenacea in the Oak Creek Canyon area in southern Coconino County, Arizona, have leaves with a narrow, lateral, undulating, purple stripe across the mid lamina. The coloration is retained even in dried specimens. In Utah, Erythranthe verbenacea is known only from the Zion Canyon area. Molecular (P. M. Beardsley et al. 2003) and morphological (G. L. Nesom 2014b) data indicate that Erythranthe verbenacea is sister to E. eastwoodiae. (Discussion copyrighted by Flora of North America; reprinted with permission.)	Erythranthe willisii is narrowly endemic over serpentine along the North Fork Feather River (including the North Branch) in Plumas County. In the original description, its range was said to include serpentine localities in closely adjacent areas of east-central Butte, Plumas, and northwestern Yuba counties, but subsequent field work has shown that these peripheral populations are E. moschata, and that E. willisii occurs only in the bottom of the Serpentine Canyon area. The most consistent and recognizable features of E. willisii are the long, sprawling stems often spread over a large area, sometimes reaching at least 45 cm and often with many crowded nodes, sessile or subsessile leaves with rounded to subcordate bases, and short pedicels, characteristically no longer than the subtending leaves (except sometimes the distal ones where subtending leaves are distinctly reduced in size). It is possible that stem growth in E. willisii is indeterminate versus determinate in E. moschata. Sessile to subsessile leaves occur in E. moschata, especially in the California Sierra Nevada, but petiole length and leaf base shape are variable within populations; lack of petioles and a rounded/subcordate base are fixed characters in E. willisii (as they are also in E. ptilota). Although large colonies of E. moschata are sometimes encountered, the individual plants tend to be erect (in California) and with few distal flowers. In the field, the dense vestiture of E. willisii is a prominent feature, but this is harder to distinguish in pressed specimens, and there is a strong tendency for purple abaxial leaf coloration in E. willisii. Phenology and flower morphology of E. willisii and E. moschata appear to be similar, but E. moschata in north-central California does not occur at as low elevations as E. willisii. (Discussion copyrighted by Flora of North America; reprinted with permission.)
Source	FNA vol. 17, p. 394.	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. 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. 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. scouleri, E. shevockii, E. sierrae, E. suksdorfii, E. taylorii, E. thermalis, E. tilingii, E. trinitiensis, E. unimaculata, E. utahensis, E. verbenacea, E. washingtonensis
Synonyms	Mimulus verbenaceus, M. cardinalis var. verbenaceus, M. lugens
Name authority	(Greene) G. L. Nesom & N. S. Fraga: Phytoneuron 2012-39: 37. (2012)	G. L. Nesom: Phytoneuron 2017-17: 7, figs. 14–22. (2017)
Web links	WildflowerSearch iNaturalist (observations) LBJ Wildflower Center SEINet Plants of the World Online Encyclopedia of Life Wikipedia Google Image Search	Local Web sites: CalFlora, CalPhotos WildflowerSearch iNaturalist (observations) LBJ Wildflower Center SEINet Plants of the World Online Encyclopedia of Life Wikipedia Google Image Search

Erythranthe willisii

crimson monkeyflower, pico Pajaro

Willis' monkeyflower

Habit

Perennials, rhizomatous.

Perennials, rhizomatous, rarely rooting at proximal nodes, usually forming large colonies, rhizomes white, usually highly branching.

Stems

erect to decumbent, usually simple, weakly 4-angled, 20–60 cm, ± glandular-villous.

usually sprawling-decumbent, branched, sometimes simple, 7–45 cm, nodes (2–)4–15+, densely glandular-villous, hairs 1–2 mm, glandular, internodes evident.

Leaves

cauline;

petiole 0 mm;

blade palmately 3–5-veined, elliptic to obovate, rhombic-ovate, or broadly spatulate, 50–75 × 15–26(–30) mm, base subcordate, subclasping, margins coarsely serrate, sometimes only distally, apex acute to obtuse, surfaces ± glandular-villous.

usually cauline, basal not persistent, distinctly separated;

petiole 0 mm, sometimes 1–2 mm at proximal nodes;

blade bicolored, purplish abaxially, pinnately veined, ovate to elliptic-ovate, midcauline 10–35 × 6–18 mm, base rounded to subcordate, margins coarsely serrate-dentate to denticulate or subentire, apex short-attenuate to acute, obtuse, or rounded, surfaces densely glandular-villous, hairs 1–2 mm, gland-tipped.

Flowers

herkogamous, 2–12, axillary at leafy medial to distal nodes.

herkogamous, (4–)8–30+, from medial to distal nodes, sometimes from all nodes.

Styles

glabrous.

Corollas

crimson, often yellow-tinged, palate ridges dark red, bilaterally symmetric, strongly bilabiate;

tube-throat tubular, 25–35 mm, exserted 13–25 mm beyond calyx margin;

abaxial limb spreading, adaxial erect, lobe apex truncate, often emarginate, throat open, palate ridges densely short-villous.

yellow, throat, tube, and proximal portion of abaxial 3 lobes with fine, red to brownish lines, weakly bilaterally or nearly radially symmetric, weakly bilabiate or nearly regular;

tube-throat narrowly funnelform, 12–15 mm, exserted beyond calyx margin;

limb 9–12 mm wide (pressed), lobes oblong-obovate, apex rounded to notched.

Fruiting pedicels

45–90(–150) mm.

4–20(–25) mm, densely glandular-villous, hairs 1–2 mm, gland-tipped.

Fruiting calyces

campanulate, weakly inflated, 20–28 mm, sparsely glandular-villosulous to stipitate-glandular, lobes triangular to ovate-triangular, apex linear-triangular.

ridge- to wing-angled, campanulate to cylindric-campanulate, weakly inflated, 7–10 mm, densely glandular-villous, lobes erect to slightly spreading, unequal, triangular to linear-lanceolate, 2–4 mm, apex acuminate-apiculate.

Capsules

included, 15–22 mm.

included, 4–5 mm.

Anthers

exserted, white-villous, thecae reflexed 45º.

included, glabrous or finely hirtellous to scabrous.

= 16.

Erythranthe verbenacea

Erythranthe willisii

Phenology

Flowering Jun–Sep.

Flowering May–Sep.

Habitat

Stream edges and beds, flood plains, around seeps and springs, canyon bottoms, moist cliff crevices and ledges.

Seepage, drainage margins, moist soils, talus, cracks and crevices, soils deprived from serpentine.

Elevation

300–2600 m. (1000–8500 ft.)

(500–)700–900 m. ((1600–)2300–3000 ft.)

Distribution

AZ; UT; Mexico (Baja California, Chihuahua, Durango, Sinaloa, Sonora)

[WildflowerSearch map]

Discussion

R. K. Vickery (1992) noted that yellow-flowered morphs of Erythranthe verbenacea occur in a population at Vasey’s Paradise in the Grand Canyon (Coconino County), 32 miles downstream from Lees Ferry.

Populations of Erythranthe verbenacea in the Oak Creek Canyon area in southern Coconino County, Arizona, have leaves with a narrow, lateral, undulating, purple stripe across the mid lamina. The coloration is retained even in dried specimens.

In Utah, Erythranthe verbenacea is known only from the Zion Canyon area.

Molecular (P. M. Beardsley et al. 2003) and morphological (G. L. Nesom 2014b) data indicate that Erythranthe verbenacea is sister to E. eastwoodiae.

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

Erythranthe willisii is narrowly endemic over serpentine along the North Fork Feather River (including the North Branch) in Plumas County. In the original description, its range was said to include serpentine localities in closely adjacent areas of east-central Butte, Plumas, and northwestern Yuba counties, but subsequent field work has shown that these peripheral populations are E. moschata, and that E. willisii occurs only in the bottom of the Serpentine Canyon area. The most consistent and recognizable features of E. willisii are the long, sprawling stems often spread over a large area, sometimes reaching at least 45 cm and often with many crowded nodes, sessile or subsessile leaves with rounded to subcordate bases, and short pedicels, characteristically no longer than the subtending leaves (except sometimes the distal ones where subtending leaves are distinctly reduced in size). It is possible that stem growth in E. willisii is indeterminate versus determinate in E. moschata. Sessile to subsessile leaves occur in E. moschata, especially in the California Sierra Nevada, but petiole length and leaf base shape are variable within populations; lack of petioles and a rounded/subcordate base are fixed characters in E. willisii (as they are also in E. ptilota). Although large colonies of E. moschata are sometimes encountered, the individual plants tend to be erect (in California) and with few distal flowers. In the field, the dense vestiture of E. willisii is a prominent feature, but this is harder to distinguish in pressed specimens, and there is a strong tendency for purple abaxial leaf coloration in E. willisii. Phenology and flower morphology of E. willisii and E. moschata appear to be similar, but E. moschata in north-central California does not occur at as low elevations as E. willisii.

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

Source

FNA vol. 17, p. 394.

FNA vol. 17, p. 401.

Parent taxa

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. 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. washingtonensis, E. willisii

Synonyms

Mimulus verbenaceus, M. cardinalis var. verbenaceus, M. lugens

Name authority

(Greene) G. L. Nesom & N. S. Fraga: Phytoneuron 2012-39: 37. (2012)

G. L. Nesom: Phytoneuron 2017-17: 7, figs. 14–22. (2017)

Web links

Unless otherwise noted, all data comes from the online version of the Flora of North America, who have made most of their content available via a Creative Commons license (CC BY 4.0).

Choose a data source: Flora of North America

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Erythranthe verbenacea

Erythranthe willisii

Erythranthe verbenacea

Erythranthe willisii