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harlequin monkeyflower, Whitney's monkeyflower

lop-seed family, monkeyflower family

Habit Herbs, annual, herbage usually drying dark. Herbs, subshrubs, or shrubs, annual or perennial, aquatic or terrestrial, sometimes fleshy, autotrophic.
Stems

erect or ascending, 10–140(–220) mm, glandular-puberulent to glandular-pubescent.

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

Leaves

usually cauline, relatively even-sized;

petiole absent;

blade narrowly elliptic to narrowly oblanceolate, linear-lanceolate, or linear, (4–)7–23(–34) × 1–5(–8) mm, margins entire, plane, not ciliate, apex acute, surfaces glandular-puberulent.

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).

Pedicels

1–3(–4) mm in fruit.

Flowers

2 per node, or 1 or 2 per node on 1 plant, chasmogamous.

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

glandular-puberulent.

Corollas

(a) magenta, darkening toward mouth and within tube, often nearly obscuring dark longitudinal stripes within, throat floor or at least palate ridges yellow, or (b) yellow, usually with maroon stripes in throat extending onto lobe bases, sometimes with maroon blotches on adaxial lateral walls of throat, colored palate ridges ending in throat, tube-throat (10–)13–18(–20) mm, limb 10–19 mm diam., not bilabiate.

Calyces

symmetrically attached to pedicels, not inflated in fruit, (3–)4–8(–10) mm, glandular-puberulent, tube not strongly plicate, lobes triangular, subequal, apex acute-apiculate, ribs narrow, darkened, purplish, thin, not strongly raised, intercostal areas pale green.

Fruits

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

Capsules

(4.5–)6–10(–13) mm.

Seeds

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

embryo straight, endosperm sparse.

Anthers

included, ciliate.

Stigmas

included, lobes equal.

2n

= 16.

Diplacus bicolor

Phrymaceae

Phenology Flowering May–Sep.
Habitat Disturbed areas, water runoff areas, granitic soils, edges of granite outcrops.
Elevation (1200–)1500–3300 m. ((3900–)4900–10800 ft.)
Distribution
from FNA
CA
[BONAP county 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

Diplacus bicolor is endemic to Fresno, Kern, and Tulare counties in the southern Sierra Nevada. The combination D. whitneyi (A. Gray) G. L. Nesom is illegitimate, as Mimulus whitneyi A. Gray 1886, is a replacement name based on Eunanus bicolor A. Gray 1868.

Diplacus bicolor, D. bigelovii, D. constrictus, D. graniticola, D. layneae, and D. thompsonii appear to be closely related species, sometimes intergrading where sympatric. Plants of each species often produce flowers at all nodes and have dark magenta corollas with nearly regular to weakly bilabiate limbs and villous vestiture. Specimens of each species dry to a dark color.

(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. 437. FNA vol. 17, p. 365. Authors: Richard K. Rabeler, Craig C. Freeman, Wayne J. Elisens.
Parent taxa Phrymaceae > Diplacus
Sibling taxa
D. angustatus, D. aridus, D. aurantiacus, D. bigelovii, D. bolanderi, D. brandegeei, D. brevipes, D. calycinus, D. cascadensis, D. clevelandii, D. clivicola, D. compactus, D. congdonii, D. constrictus, D. cusickii, D. cusickioides, D. deschutesensis, D. douglasii, D. fremontii, D. grandiflorus, D. graniticola, D. jepsonii, D. johnstonii, D. kelloggii, D. layneae, D. leptaleus, D. linearis, D. longiflorus, D. mephiticus, D. mohavensis, D. nanus, D. ovatus, D. parryi, D. parviflorus, D. pictus, D. pulchellus, D. puniceus, D. pygmaeus, D. rattanii, D. rupicola, D. rutilus, D. thompsonii, D. torreyi, D. traskiae, D. tricolor, D. vandenbergensis, D. viscidus
Subordinate taxa
Diplacus, Erythranthe, Glossostigma, Mimetanthe, Mimulus, Phryma
Synonyms Eunanus bicolor, M. nanus var. bicolor, M. whitneyi
Name authority (A. Gray) Hrusa: Phytoneuron 2014-17: 1. (2014) Schauer
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