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crimson monkeyflower, pico Pajaro

lop-seed family, monkeyflower family

Habit Perennials, rhizomatous. Herbs, subshrubs, or shrubs, annual or perennial, aquatic or terrestrial, sometimes fleshy, autotrophic.
Stems

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

erect or ascending to prostrate, 4-angled, sometimes winged.

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.

deciduous or persistent, basal and cauline or all cauline, rarely subrosulate or rosulate (Erythranthe), opposite, or alternate distally, simple;

stipules absent;

petiole present or absent;

blade fleshy, semi-fleshy, or not, not leathery, margins entire or toothed.

Inflorescences

terminal and axillary racemes or flowers solitary (Glossostigma, some Erythranthe, some annual plants);

flowers erect to nodding or strongly reflexed and appressed to inflorescence axis (Phryma).

Flowers

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

bisexual, perianth and androecium hypogynous;

sepals (3 or)4 or 5, proximally connate, calyx radially or bilaterally symmetric;

petals 3–5, proximally connate, corolla bilaterally symmetric, rarely nearly radially in reduced forms, strongly to weakly bilabiate, rarely nearly regular, salverform to tubular-funnelform, funnelform, campanulate, or compressed;

stamens (2–)4, adnate to corolla, didynamous [both pairs of equal length in autogamous forms], staminode 0;

pistil 1, 2-carpellate, ovary superior, (1 or)2-locular, placentation axile, basal (Phryma), or parietal (Diplacus, Mimetanthe);

ovules anatropous or orthotropous (Phryma), unitegmic, tenuinucellate;

style 1;

stigma 1, 2-lobed.

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.

Fruiting pedicels

45–90(–150) mm.

Fruiting calyces

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

Fruits

capsules, dehiscence loculicidal [septicidal or irregular], or achenes [berry].

Capsules

included, 15–22 mm.

Seeds

1–2000, yellowish brown or brown, narrowly ellipsoid, slightly flattened bilaterally;

embryo straight, endosperm sparse.

Anthers

exserted, white-villous, thecae reflexed 45º.

2n

= 16.

Erythranthe verbenacea

Phrymaceae

Phenology Flowering Jun–Sep.
Habitat Stream edges and beds, flood plains, around seeps and springs, canyon bottoms, moist cliff crevices and ledges.
Elevation 300–2600 m. (1000–8500 ft.)
Distribution
from FNA
AZ; UT; Mexico (Baja California, Chihuahua, Durango, Sinaloa, Sonora)
[WildflowerSearch map]
North America; Mexico; Central America; w South America (primarily Andean); s Asia (India); se Asia; e Africa; Indian Ocean Islands (Madagascar); Pacific Islands (New Zealand); Australia [Introduced in Europe, s Africa]
[BONAP county 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.)

Genera 13, species ca. 200 (6 genera, 139 species in the flora).

Over one half of the species in Phrymaceae are members of Diplacus and Erythranthe and together include over 160 species; all other genera each have seven or fewer species.

Until recently, Phrymaceae consisted only of Phryma leptostachya, a taxonomically isolated species of eastern North America and eastern Asia. Molecular studies have established a relationship not with the Verbenaceae, as was earlier postulated (see H. L. Whipple 1972; R. Venkata Ramana et al. 2000), but rather with Mimulus and other genera, suggesting that Phrymaceae should be enlarged. The sequence of genera in Phrymaceae here follows the phylogeny proposed by P. M. Beardsley and R. G. Olmstead (2002).

One of the major lineages of Phrymaceae is primarily a Southern Hemisphere group ranging from Australia and New Zealand to southeastern and south Asia (India), Madagascar, and South Africa. Mimulus in the narrow sense, including the two endemic North American species, is part of this group, which includes 24 species in seven genera. The largest major lineage includes 158 species in five genera from North America, South America, and southeast Asia. This lineage includes two genera from Mexico and Central America: Hemichaena Bentham, which is sister to the North American Diplacus and Mimetanthe, and Leucocarpus D. Don, which is sister to the American and Asian Erythranthe.

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

Key
1. Plants aquatic or semi-aquatic, mat-forming.
→ 2
2. Stems functionally stolons, leaves on stolons; sepals 3; leaf blades fleshy.
Glossostigma
2. Stems prostrate to decumbent or erect; sepals 5, sometimes 3 in reduced forms; leaf blades sometimes thickened or semi-fleshy.
Erythranthe
1. Plants terrestrial or, if semi-aquatic, not mat-forming.
→ 3
3. Flowers strongly reflexed and appressed to inflorescence axes in fruit; fruits achenes; bracteoles present.
Phryma
3. Flowers lateral or erect to nodding, not strongly reflexed and appressed in fruit; fruits capsules; bracteoles absent.
→ 4
4. Fruit apices rounded to truncate; placentation axile.
→ 5
5. Leaf venation brochidodromous; stamens adnate to middle of corolla.
Mimulus
5. Leaf venation acrodromous (veins usually basal only, sometimes basal and suprabasal); stamens adnate proximal to middle of corolla.
Erythranthe
4. Fruit apices attenuate; placentation parietal.
→ 6
6. Pedicels ± equal to or slightly longer than calyces; calyx lobe midveins low-rounded, not wing-angled; fruit walls densely pustulate-glandular.
Mimetanthe
6. Pedicels nearly absent or shorter than calyces, rarely ± equal to or slightly longer than calyces; calyx lobe midveins angled or wing-angled; fruit walls smooth, eglandular.
Diplacus
Source FNA vol. 17, p. 394. FNA vol. 17, p. 365. Authors: Richard K. Rabeler, Craig C. Freeman, Wayne J. Elisens.
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
Subordinate taxa
Diplacus, Erythranthe, Glossostigma, Mimetanthe, Mimulus, Phryma
Synonyms Mimulus verbenaceus, M. cardinalis var. verbenaceus, M. lugens
Name authority (Greene) G. L. Nesom & N. S. Fraga: Phytoneuron 2012-39: 37. (2012) Schauer
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