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TAXONOMY OF ERYTHRANTHE SECT. MIMULOSMA (PHRYMACEAE) 

Guy L. Nesom 

2925 Hartwood Drive 

Fort Worth, Texas 76109 

guynesom@sbcglobal.net 

ABSTRACT 

Seventeen species are treated as members of Erythranthe sect. Mimuhsma in North America 
north of Mexico: E. ampliata, E. arenaria, Erythranthe austrolatidens Nesom, sp. nov. (Baja 
California Sur), E. breviflora, E. floribunda, E. geniculate! (synonym: Mimulus dudleyi), E. 
hymenophylla, E. inflatula (synonym: Mimulus evanesce/is), E. inodora. E. jungermannioides, E. 
laiidens, E. moniliformis, E. moschata, E norrisii, E. patula, .£', pulsiferae, and E. washingtonensis. 
The section also is hypothesized also to include one Russian species (E. stolonifera) for a total of 18 
species. Descriptions, typifications, a key to species, and distribution maps are provided. 

KEY WORDS: Mimulus moschatus, Mimulus floribundus, Erythranthe sect. Mimulosma, Mimulus 
moschatus alliance, Phrymaceae 



Sect Mimulosma of the genus Erythranthe (formerly part of Mimulus; Barker et al. 2012) 
includes 19 species as treated here, including the widespread E. moschata and E. floribunda, 
primarily centered in their native ranges to the western USA. The group has often been referred to as 
the "Mimulus moschatus alliance." Grant (1924) placed these species within her more broadly 

encompassing Mimulus sect. Paradanthus. 

North American species added here since study of the group by Whittall (1999), Carlson 
(2002), and Whittall et al. (2006) are Erythranthe moniliformis, E. inodora, E. arenaria, and E. 
plotocalyx. Mimulus evanescens proves to have been earlier named as M. inflatulus and an earlier 
name for Mimulus dudleyi is M. geniculatus. The group also includes one species endemic to 
southeastern Russia. 

The taxonomy of sect. Mimulosma (as "the Mimulus moschatus alliance") has been studied 
using pollen morphology (Argue 1980, 1986), vegetative and reproductive morphology (Carlson 
2002), and molecular data (Whittall 1999; Whittall et al. 2006). Molecular data show the group to be 
distinct, but Carlson was unable to find any morphological synapomorphies for it and described it as a 
"morphologically cryptic clade." Within Erythranthe, species of the group are generally recognized 
by their herbaceous habit, yellow corollas, equal to subequal calyx lobes, and glandular vestiture. 
Carlson (2002) noted that "traits ... associated with most ingroup species include a tetraploid 
chromosome number (N = 16), progressively bifurcated stem architecture, viscid herbage, anthers 
that open completely, and unequal theca." 

Most resolutions of molecular and morphological phylogenies divide the species of 

Erythranthe sect Mimulosma between two main groups — a northern group (the "Columbia River 
clade"; see Maps 1 and 2) and a more southern group (the "Sierra Nevada clade). The latter includes 
E. floribunda, E. geniculate!., E. norrisii, E. arenaria, E. moschata, E. inodora, and E. moniliformis. 
The Columbia River clade includes E. ampliata, E. patula, E. hymenophylla, E. Jungermannioides, E. 
washingtonensis, and E. breviflora. Erythranthe ampliata, E. patula, and E. hymenophylla are a 
subgroup, the "Snake River clade," within the northern group. The positions of E. laiidens and E. 
pulsiferae are equivocal; E, plotocalyx, first described here, appears to be similar to E. latidens. 



Nesom: Taxonomy of Erythranthes 



Erythranthe inflatula is hypothesized to he of hybrid origin between E. latidens and E. breviflora. 
The Russian E. stolonifera apparently is most closely related to E. moschata-moniliformis-inodora. 
Species not included in earlier studies are interpolated here on the basis of morphology and 
geography. 

This account is based primarily on study of collections from ARIZ, BRIT-SMU-VDB, DAV, 
MO, ND-Greene, NMC, PH, SRSC, TEX-LL, UC-JEPS, and UT. Specimens are cited only for some 
of the species or variants that have not been generally recognized, but collections at herbaria above 
have been annotated in documentation of the concepts. 

ERYTHRANTHE sect. Mimulosma Nesom & Fraga, Phytoneuron 2012-n: 0. 2012. 

TYPE; Erythranthe moschata (Douglas ex Lindl.) Nesom 

Annuals and fibrous -rooted or taprooted or perennials from rhizomes; vestiture of gland- 
tipped hairs, varying from minutely stipitate-glandular to villous -glandular, sometimes aromatic. 
Leaves pmnately or suhpinnately to palmately veined. Calyces strongly ridge-angled to wing-angled 
at maturity, lobes mostly of equal or subequal length. Corollas yellow or white to pinkish or flesh- 
colored, strongly bilabiate to weakly bilabiate or nearly regular; palate puberulent to short-villous 
with stiff, usually clavate hairs. Anthers included. Capsules included (often slightly exserted in E. 
norrisii). Base chromosome number, x = 8. 

Two pollen types were recognized by Argue (1980, 1986) among species of sect. 
Mimulosma. All pollen is tricolporate with multiple, elongate endoapertures. Those with the sexine 2 
configuration predominantly microreticulate without supramurial granules or spinules are type lib: 

Erythranthe arenaria, E. geniculata, E. floribunda, and E. moniliformis (presumably E. norrisii, 
which was not sampled, also has type lib). The others (including E. moschata}, with supramurial 
granules or spinules, are type lie. 

An autogamous mating system lias evolved independently several times within Erythranthe 
sect. Mimulosma. Floral characters associated with a shift from allogamy to autogamy include 
decreased bilateral symmetry, less reflexed upper corolla lobes, and reduced overall corolla length 
(with corresponding reduction of anther/stigma separation). Carlson (2002) and Whittall et al. (2006) 
discussed the evolution of this syndrome and observed that it characterizes E. floribunda, E. patula, 
E. breviflora, E. inflatula, and. E. latidens and it is noted here mat similar features characterize E. 
plotocalyx. 

Key to the American species 

1. Plants from rhizomes and/or stolons. 



2. Calyx lobes 1-2 mm apices i m-idtd to nmuotiaie aMh^is ^lilous shks scabrous; stolons 
forming overwintering turions; plants characteristically of cliff faces. 

1. Erythranthe j ungermannioides 

2. Calyx lobes 2-9 mm, apices acute or acuminate; anthers glabrous or pubescent; styles glabrous; 
stolons without turions; plants usually of habitats other than cliff faces. 

3. Stems erect to ascending-erect; leaves often congested 10. Erythranthe moniliformis 

3. Stems procumbent to decumbent or decumbent-ascending; leaves distinctly separated. 



Nesom: Taxonomy of Erythranthe s 



4. Cauline leaves all petiolate, blades (10-)15^IO(-50); fruiting pedicels (7-)10-25(-40) 
mm; calyx lobes triangular to linear-lanceolate or narrow h ijnngulai ■ummnjte 2 ~i mm 
corolla tube-throats 11-16 mm; anthers glabrous to subglabrous . 9. Erythranthe moschata 
4. Cauline leaves usually sessile (proximal sometimes short-petiolate or subpetiolate), blades 
generally oblong-lanceolate, 30-70 mm; fruiting pedicels (15-)22-50 mm; calyx lobes linear- 
lanceolate to narrowly triangular with linear-acuminate apices, 5-9 mm; corolla tube-throats 
15-18 mm; anthers strongly to weakly hirsute-hirtellous 11. Erythranthe inodora 

Plants fibrous-rooted or taprooted, without rhizomes or stolons. 

5. Cauline leaves gradually petiolate to sessile or subsessile, blades generally elliptic to lanceolate- 
elliptic or ovate-elliptic with an attenuate base, palmately 3-5-veined. 

6. Stems and pedicels villous-glandular with gland-tipped hairs 0.2— 1(— 1.5) mm. 

7. Corolla tube-throats 9-12(-14) mm, red spots of lower lip conspicuous; calyx prominently 

red-dotted 13. Erythranthe arenaria 

7. Corolla tube-throats 5-6 mm, red spots of lower lip small and indistinct; calyx commonly 
without red dots 

8. All leaves petiolate; erect to decumbent, sometimes procumbent-trailing, branching at 

proximal to distal nodes 14. Erythranthe floribunda (in part, in Arizona) 

8. Medial to distal cauline leaves sessile; stems erect to erect-ascending, mostly branched 
at the base 15. Erythranthe austrolatidens 

6. Stems and pedicels subglabrous ro ws*de-glanduLt ot rmnutely stipitate-glandular with 
gland-tipped hairs 0. 1-0.3 mm, without villous hairs. 

9. Petioles 2-9 mm, distinctly 3-nerved (winged); fruiting pedicels 12-38 mm, divergent- 
arcuate 12. Erythranthe pulsiferae 

9. Petioles absent or 1-3 mm, 1-nerved; fruiting pedicels 5-28 mm, straight. 

10. Fruiting calyces minutely stipitate-glandular, 8-12 mm; fruiting pedicels 11-28 mm; 

corolla tube-throats 5-6(-8) mm; leaves basal and cauline 8. Erythranthe latidens 

10. Fruiting calyces sparsely and minutely hirtellous, eglandular or sparsely sessile- 
glandular, 5-6 mm or 7-11 mm; fruiting pedicels 5-11 mm or 7-18 mm; leaves mostly 
cauline (basal deciduous by flowering). 

11. Fruiting calyces 5-6 mm; fruiting pedicels 5-11 mm; corolla tube-throats 3.5-5 
mm, not exserted beyond the calyx margin; all leaves short-petiolate 

6. Erythranthe breviflora 

11. Fruiting calyces 7-11 mm; fruiting pedicels 7-18 mm; corolla tube-throats 5-6(- 
8) mm, exserted 1-3 mm beyond calyx margin; midcauline and distal leaves 
subpetiolate to subsessile 7. Erythranthe inflatula 

5, Cauline leaves abruptly and distinctly petiolate, blades generally ovate with a rounded to 
truncate or cordate base; palmately or pinnately veined. 

12. Stems prostrate to ascending-erect, sharply bent at the basal nodes; fruiting pedicels 
divergent at ca. right angles from the stem, often closely paired; calyx lobes ovate-rounded 
3. Erythranthe hymenophylla 



Nesom: Taxonomy of Erythranthe s 



\2. Stems erect to prostrate, decumbent, or ascending, straight (if erect) or geniculate at nodes; 
fruiting pedicels suberect to ascending-erect, not paired; calyx lobes acute to deltate or 
shallowly deltate-subulate. 

13. Stems and pedicels stipitate-glanduiar (gland-tipped trichomes unicellular, 0.05-0.2 
mm long); leaf blades palmately veined. 

14. Corollas regular to weakly bilabiate, tube-throats 7-8 mm 4. Erythranthe patula 
14. Corollas strongly bilabiate, tube-throats 8-12(-14)mm. 

15. Stems, leaves, and calyces villous-glandular with vitreous, flattened, 
multicellular, gland-tipped hairs 0.1-0.8 mm long, leaves densely hairy; stems 

terete; styles hispid-hirtellous 2. Erythranthe washingtonensis 

15. Stems, leaves, and calyces minutely sessile to subsessile glandular with gland- 
tipped hairs 0.05-0.2 mm long, leaves very sparsely glandular; styles glabrous; 
stems 4-angled 5. Erythranthe ampliata 

13. Stems and pedicels villous-glandular (gland-tipped trichomes multicellular, mostly 0.5- 
1.2(-2) mm long); leaf blades pinnately to subpinnately veined. 

16. Plants mostly erect to ascending-erect; calyces greenish; styles hispid-hirtellous; 

Oregon and Washington 2. Erythranthe washingtonensis 

16. Plants erect to prostrate, decumbent, or ascending; calyces commonly red-spotted; 

styles glabrous; widespread (15) or California (16, 17). 

17. Corolla tube-throats (4-)5-10 mm, limbs expanded 3-4 mm across (pressed) 

14. Erythranthe floribunda 

17. Corolla tube-throats 9-11 mm or 12-16 mm, limbs expanded 8-16 mm across 
(pressed). 

18. Calyces with shallowly wing-an°led ribs, lobe<. triangular-acute i-nd erect 
to spreading or recurving-spreading in fruit; corollas without white patches, 
tube-throats 9-12 mm, expanded limb ca. 10-18 mm across 

16. Erythranthe geniculata 

18. Calyces with rounded-thickened ribs, lobes linear-oblong and incurved in 
fruit; corollas with white patches on the lower lip, tube-throats 12-16 mm, 
expanded limb 15-30 mm across 17. Erythranthe norrisii 

1. Erythranthe jungermannioides (Suksdorf) Nesom, Phytoneuron 2012-39: 38. 2012. Mimulus 
jungermannioides Suksdorf, Deutsche Bot. Monatsschr. 18: 154. 1900. TYPE: USA. 
Washington. Klickitat Co.: steep, overhanging, damp cliffs near Bingen, Aug-Nov 1892, 
W.N. Suksdorf 1470 (holotype: MO!; isotypes: F, NY digital image!, UC!, US digital image!, 
WTU). 

Perennial, from thin, above-ground stolons or runners that form terminal bud-like structures 
(overwintering turions). Stems decumbent to procumbent, 5-38(-60) cm, simple or branching near 
the base. Stems and pedicels densely glandular-villous with gland-tipped hairs 0.5-1. 2(-l. 5) mm. 
Leaves cauline, basal not persistent blades broadly ovate to broadly lanceolate, 7-35(-40) mm x 8- 
25 mm, subpahnately to pinnately veined, margins sharply and irregularly dentate to denticulate, apex 
acute to obtuse, base rounded; petioles mostly 2-5(-20) mm. Fruiting calyces cylindric-urceolate, 
6-12 mm, plicate-angled, weakly inflated, glandular-villous, lobes subequal, 1-1.5 mm, rounded- 



Nesom: Taxonomy of Erythranthe s 



, sobered, ciliate. Fruiting pedicels 15-35 mm. Corollas yellow with scattered red spots 
and two white patches at the tips of the palate ridges, strongly bilabiate, tube-throats funnelform, 
(12-)16-20(-24) mm, limbs expanded 8-10 mm (pressed), lobes obovate-oblong, apex rounded to 
truncate. Styles scabrous. Anthers glabrous. Capsules elliptic-lanceolate, 5-9 mm. In = 32. Map 
1. 

Flowering May-Jul(-Aug). Basalt crevices in seepage zones in vertical cliff faces and 
canyon wails, within xeric sagebrush communities; 100-400(-1200) m; Oreg., Wash. The record on 
Map 1 for Jefferson Co., Oregon, is added from OBIC (20 11). The Klickitat Co., Washington, 
occurrence is termed "historical" by WNHP (2005), which also notes that "A sighting of the plant on 
the WA side was made in the early 1990s, but the location and presence of the species need to be 
confirmed." 

Molecular data place Erythranthe jungermannioides and E. washingtonensis as sister species. 

2. Erythranthe washingtonensis (Gandoger) Nesom, Phytoneuron 2012-39: 39. 2012. Mimulus 
washingtonensis Gandoger, Bull. Soc. Bot. France 66: 218. 1919. TYPE: USA. Washington. 
Klickitat Co.: low sandy banks of the Columbia, Bingen, Oct-Nov 1885, W.N. Suksdorf 560 
(holotype: LY; isotypes: CAS digital image!, CU, DS digital image!, GH, MO 2 sheets!, ND- 
Greene!, NY 2 sheets digital images!, ORE digital image!. OS digital image!, PH!, UC 2 
sheets!, US digital image!, WS, WTU). 

Printed labels by Meinke in 1987 on type specimens give this: "ISOTYPE of 
Mimulus washingtonensis Gandoger, according to number only; ... The numerous sheets 
distributed under no 560 collectively represent a hybrid swarm of undetermined parentage, 
probably involving M. washingtonensis, M. patulus, and M. floribundus." Meinke did not 
provide document to substantiate his observation. Among the plants I have studied of the 
type collection, density of villous hairs near the stem bases and on the leaf blades varies, and 
one of the 16 individual plants on the two UC sheets has minutely hirtellous lower cauiine 
vestiture, but all plants I have seen of the type collection have villous-glandular leaf surfaces, 
which distinguishes them from E. ampliata. Matt Carlson (pers. comm.) also notes that he 
finds the vestiture of the type collection uncharacteristic of the species in its wider 
occurrence, but whether this has resulted from hybridization is not evident. 

Annuals, fibrous-rooted or filiform-taprooted. Stems erect to ascending, 5-25 cm, often 
many-branched, terete. Stems and pedicels puberulent-glandular to villous-glandular with gland- 
tipped hairs 0.1-0.8 mm, hairs sometimes vitreous, flattened, and distinctly multicellular. Leaves 
cauiine, basal not persistent, blades deltate or ovate to ovate-lanceolate, 4-16( -23) mm x 2-ll(-16) 
mm, palmately veined, margins denticulate to entire, apex acute, base rounded to cuneate or truncate, 
petioles 2-14 mm. Fruiting calyces tubular, 6-8 mm, ridge-angled, weakly inflated, densely and 
minutely stipitate-glandular, lobes subequal, 0.8-1.2 mm, shallowly deltate. suberect, ciliate. 
Fruiting pedicels 20-50 mm, divergent at nearly right angles. Corollas yellow with small reddish 
brown dots and two white patches on the lower lip, strongly bilabiate, tube-throats funnelform, 8-10 
mm, limbs spreading 7-10 mm (pressed), lobes obovate-oblong, apex rounded to rounded-cuneate. 
Styles hispid-hirtellous. Anthers glabrous. Capsules ellipsoid to ellipsoid-fusiform, 5-8.5 mm. 2n 
= 32. Map 1. 

Flowering May-Sep. Shallow basalt gravels in narrow channels and intermittent streams, 
sandy stream banks, open slopes, rocky shelves near seeps; 700-1300 m; Oreg., Wash. 

Despite the implication of the epithet, except for the type collection Erythranthe 
washingtonensis occurs most abundantly in the John Day River drainage of eastern Oregon. 



Nesom: Taxonomy of Erythranthe s 



3. Erythranthe liymenophylla (Meinke) Nesom, Phytoneuron 2012-39: 38. 2012. Mimulus 
hymenophyllus Meinke, Madrono 30: 147, plate 1. 1983. TYPE: USA. Oregon. Wallowa Co.: E side 
of Horse Creek, ca. 12 km S of the Imnaha River and ca. 21 km W of the Snake River, 1075 m, 2 Jul 
1980, R.J. Meinke & Kennison 2656 (holotype: OS digital image!; isotypes: ID, N\ digital image! 
ORE digital image!, UC!, US digital image!, WS, WTU). 

Annual, filiform-taprooted. Stems prostrate to ascending-erect, 5-25 cm, simple or few- 
branched. Stems and pedicels glandular-puberulent to glandular-villous with vitreous, flattened, 
multicellular, gland-tipped hairs 0. 1-0.8 mm, ail hairs glandular. Leaves basal and cauiine, largest at 
midstem, blades broadly lanceolate to ovate, 10-35 mm x 10-30 mm, pinnately veined, distinctly 
membranous, margins coarsely dentate to shallowly denticulate or entire, apex acute to obtuse, base 
cuneate to shallowly cordate, petioles 6-30 mm. Fruiting calyces tubular-campanulate, angled, 5-7 
mm, slightly inflated, sparse!} ^tipita'e-olandulai bbes subequal, 0.5-1.2 mm, ovate-rounded, 
suberect, ciliate. Fruiting pedicels 10-45 mm, negatively phototropic, causing the capsules to be 
pressed against the cliff face or into a crevice by the time of dehiscence. Corollas light yellow with 
red or purple spots on the throat and lower lobes, sometimes with small white patches, weakly 
bilabiate, tube-throats funnelform, 10-14 mm, lobes obovate-oblong, apex rounded to truncate or 
notched. Styles glabrous. Anthers glabrous. Capsules ovoid, 3-6 mm. 2n = 32. Map 1. 

Flowering Apr-Aug(-Sep). Steep, seasonally moist basalt cliffs with W or SW exposure, 
within mesic coniferous forests; 800-1300 m; Idaho, Mont., Oreg. 

Until recently, Erythranthe hymenophytta has been thought to be restricted to deep canyons 
of Horse Creek and Cow Creek in southeast Wallowa County, Oregon. The Oregon Biodiversity 
Information Center (2010) notes that the species also is known in Idaho and Montana — county 
records are added from BONAP (20 1 1). 

Meinke (1983) observed that plants of E. hymenophytta have reflexed ("strongly negatively 
phototropic") fruiting pedicels that increase seed dispersal back onto the vertical cliff wall, the 
characteristic habitat of the species. The "hanging" habit of E. hymenophytta is reflected in a shasp 
(90°-180°) bend in the basal nodes and the long pedicels that are closely paired and divergent in 

parallel at about right angles from the stem. The species also is characterized by it very short calyx to 
corolla length, relatively short capsules, and very large seeds. 

4. Erythranthe patula (Pennell) Nesom, Phytoneuron 2012-39: 39. 2012. Mimulus patutus Pennell, 

Proc. Acad. Natl Sci. Philadelphia 99: 162. 1947. TYPE: USA. Washington. Whitman Co.: 

Wawawai, along irrigation ditches, May 1897, A.D.E. Elmer 752 (holotype: PH!; isotypes: 

MO-2 sheets!, SMU!, US digital image!). 

Annuals, fibrous-rooted or filiform-taprooted. Stems erect to ascending, (3-)5-15(-24) cm, 
usually simple. Stems and pedicels stipitate-glandular with gland-tipped hairs 0.2-0.5 mm. Leaves 
cauiine, basal not persistent, blades deitate or ovate to ovate-lanceolate, 4-12(-17) mm x 3-10(-14) 
mm, palmately 3-veined, margins usually denticulate, apex acute to obtuse, base rounded to cuneate- 
truncate, petioles (5-)8-25 mm. Fruiting calyces tubular, 5-6(-7) mm, not inflated or weakly so, 
sparsely stipitate-glandular to sparsely hirtellous, lobes subequal, 0.7-1.1 mm, deitate, suberect, 
ciliate. Fruiting pedicels 10-25(-38) mm. Corollas yellow, usually with a few red or brownish dots 
on lower lip, regular to very weakly bilabiate, tube-throats funnelform, 7-8 mm, lobes oblong, apex 
rounded to truncate. Styles glabrous. Anthers glabrous. Capsules ellipsoid to narrowly obovoid, 4- 
6 mm. 2n = 32. Map 1. 

Flowering Apr-May(-Jul, -Aug). Ephemeral seqis, springs, rocky stream banks, moist 
basalt, fine gravel on top of bedrock, muddy hillside seeps, crevices: 200-1900 m (-2900 m in 
Montana and Wyoming); Alberta; Idaho, Mont., Oreg., Wash., Wyo. 



Nesom: Taxonomy of Erythranthe s 



Erythranthe patula i^ djstmcU\e in us long-petiolate leaves with < 
weakly bilabiate to nearly regular corollas. 



In his dissertation, Meinke (1992) proposed to recognize Mimulus patulus "var. montanus," 
characterized by a cauline vestiture of a mixture of shptt.itc-Uandulai h'irs and minute (0.1-0.2 mm), 
sharp-pointed eglandular hairs and with a geographic distribution considerably beyond the typical 
expression in its range of two or three counties at the corner of eastern Oregon and Washington. The 
vestiture of typical Mimulm patulus is constituted only by minute, stipitate-glandular hairs. "Var. 
montanus" was not mentioned again in later publications of which Meinke was author or coauthor 
(Meinke 1995; Whittall et al. 2006) and the MO specimen cited as an isotype of "var. 
cannot be located at MO or UC. 



In any case, intergrades between the two vestiture types appear to indicate that the \ 
variable within a single species. Meinke (1992) noted that a collection from Wallowa Co., Oregon 
(Peck 18282, NY, WILLU) included plants of both var. patulus and var. montanus. A collection 
from Idaho Co., Idaho (Gray 5571, MO), might also be interpreted as a mixed collection in the same 
way. Plants of other collections in 2009 by Karen Gray (MO) from Nez Perce and Idaho counties, 
Idaho, have stipitate-glandular pedicels but hirtellous, eglandular stems. Collections from Teton Co., 
Wyoming, have stipitate-glandular stems and pedicels but hirtellous calyces (Payson & Payson 2226; 
Williams 875; Williams 992; all MO) and would strictly be identified as var. patula. Anderson 366 
(UC) from Teton Co. has sparsely glandular calyces. Analogous variation in vestiture occurs among 
populations of Erythranthe breviflora as well as in E. guttata (Fischer ex DC.) Nesom of sect. 
Simiolus (Nesom 2012). 

5. Erythranthe ampliata (AL. Grant) Nesom, Phytoneuron 2012-39: 38. 2012. Mimulus ampliatus 
A. L. Grant, Ann. Missouri Bot. Gard. 11: 214. 1924. TYPE: USA. Idaho. Nez Perce Co.: 
about Lake Waha, 2000-3500 ft, 27 Jun 1896, A.A. and E.G. Heller 3330 (holotype: MO!; 
isotypes: JEPS, MO!, ND-Greene!, UC). The original labels were distributed with a 
handwritten identification of "Mimulus longipedunculatus Heller, n. sp.," but that name was 
never published. 

Annuals, fibrous-rooted or filiform-taprooted. Stems erect to ascending, 5-17 cm, often 
many-branched, 4-angled. Stems and pedicels sessile- to subsessile-glandular with gland-tipped 
hairs 0.05-0.2 mm. Leaves cauline. basal not persistent, blades broadly ovate to lanceolate, 8-25 
mm x 5-19 mm, palmately veined, margins dentate to coarsely denticulate, apex acute to obtuse, base 
cuneate, petioles 8-20 mm. Fruiting cdhcts tubulai-camp^nulate. 6-8 mm, not inflated or weakly 
so, minutely sparsely stipitate-glandular to glabrous, lobes subequal, triangular-acute, suberect, 
ciliate. Fruiting pedicels 10-22 mm. Corollas deep yellow with a few brownish dots on the lower 
lip, sometimes with small white patches, strongly bilabiate, tube-throats broadly funnelform, 8-12(- 
14) mm, lobes obovate-oblong, apex rounded to truncate. Styles glabrous. Anthers glabrous. 
Capsules fusiform to narrowly ellipsoid, 5-6 mm. Chromosome number unknown. Map 1. 

Flowering Jun-Jul. Basalt outcrops, seepy roadcuts, grassland seeps; 900-1700 m; Idaho 
(Nez Perce and Idaho counties). 

Meinke referred to Mimulus ampliatus as a variety (1992) or subspecies (1995) of M. 
washingtonensis, but he apparently never made the formal combination. As understood here, 
Erythranthe ampliata and E. washingtonensis have disjunct geographic distributions and are non- 
intergrading in morphology (as in the key). The geography and vestiture of E. ampliata are more 
similar to E. patula than to E washingtonensis. 



Nesom: Taxonomy of Erythranthe s 



6. Erythranthe breviflora (Piper) Nesom, Phytoneuron 2012-39: 38. 2012. Mimulus breviflorus 

Piper, Bull. Torrey Bot. Club 28: 45. 1901. TYPE: USA. Washington. Whitman Co.: 
Pullman, 3 Jul 1894, C.V. Piper 1858 (holotype: WS; isotypes: OS digital image!, UC, US 
digital image!). 

Annuals, shallowly fibrous-rooted. Stems ascending, 4-15 cm, branched at lower and 
middle nodes. Stems and pedicels (and leaves) minutely stipitate-glanduiar with gland-tipped hairs 
0.1-0.3 mm, without villous hairs, sometimes minutely hirtellous with minute-sharp-pointed 
eglandular hairs (as in E. patuld). Leaves mostly cauline (basal mostly deciduous by flowering), 
blades narrowly ovate or narrowly lanceolate to elliptic or elliptic-lanceolate, largest mostly 5-15 mm 
x 2-6 mm, relatively even-sized, or slightly reduced distally, palmately 3-veined, margins entire to 
mucronulate or denticulate, apex acute to obtuse, base attenuate, narrowed to short (1-3 mm) petiolar 
regions or subpetiolate to subsessile. Fruiting calyces campanulate becoming ovoid-ellipsoid to 
campanulate, 5-6 mm x 3.5-6 mm, winged and plicate-angled, distinctly inflated, sparsely and 
minutely hirtellous, eglandular or sometimes sparsely sessile-glandular, lobes subequal, 0.5-1 mm, 
ovate-deltate, suberect, ciliate. Fruiting pedicels 5-11 mm, straight. Corollas yellow, spotted or 
striped, tube-throats cylindric to narrowly funnelform, 3.5-5 mm, not exserted beyond the calyx rim, 
limb weakly bilabiate, barely widened, lobes broadly obovate, apex rounded. Styles glabrous. 
Anthers glabrous. Capsules oblong-ovoid to ovoid-cylindric, 4-6 mm. Chromosome number 
unknown. Map 2. 

Flowering May-Jul. Stream and lake sides, gravel bars, springs, moist slopes, damp swales 
between dunes, along trails; 1000-2300 m; British Columbia; Calif.. Idaho, Mont, Nev., Oreg., 
Wash. 

Voucher specimens for range extremities. British Columbia. 21.5 mi by road on Rossland- 

Cascade road from junction near Rossland, occasional on rocky, grassy, S-facing slopes, 4250 ft, 5 
Aug 1953, Colder and Saville 11483 (UC). Nevada. Humboldt Co.: Santa Rosa Mts, Martin Creek, 
moist seep under Veratrum, 7500 ft, 25 Jul 1940, Munz 16152 (UC). 

7. Erythranthe inflatula (Suksdorf) Nesom, Phytoneuron 2012-39: 38. 2012. Mimulus inflatulus 

Suksdorf, Werdenda 1: 38. 1927. TYPE: USA. Washington. Klickitat Co.: In cultivation at 
Bingen, "25 May 1901—10 Jun 1903," W. Suksdorf 991 6 (possible holotype: WS; duplicates: 
MO!, UC!). MO and UC sheets were annotated as 'isolectotype' by R.J. Meinke in 1987. 

The type sheet of Mimulus inflatulus at MO was identified by annotation by R.J. 
Meinke in 1987 as Mimulus breviflorus subsp. robustus Meinke (an unpublished name). He 
annotated a collection from Harney Co., Oregon (M.E. Peck 21389, NY digital image!) as 
"isotype" of Mimulus brevifolius var. robustus. Neither' M. inflatulus nor the name "subsp. 
robustus" was mentioned in subsequent publications by Meinke (1995a, 1995b, 2007). 
Mimulus evanescens Meinke, Great Basin Naturalist 55: 250, plate 1. 1995. TYPE: USA. California. 
Lassen Co.: 20.5 kmEof Adin, N side of Ash Valley Rd.. caO.l kmE of the Lassen National 
Forest boundary, in broken boulders and heavy gravel abutting Moll Reservoir, ca 1500 m, 27 
Jun 1990, RJ. Meinke & T. Kaye 5900 (holotype: OS; isotypes: MO, NY, RM UC, US, 
UTC). Deposition of types is as cited by Meinke but judging from their absence on online 
databases associated with those herbaria, it would seem that none of the specimens have been 
distributed. Close study of North American Mimulus sensu lato at MO and UC did not bring 
a duplicate of 5900 to light. A topotype has been studied (Schoenig 98- 70), as cited below). 

Annuals, fibrous-rooted or filiform-taprooted. Stems erect to ascending, 6-20(-25) cm, 
simple or branched at lower and middle nodes. Stems and pedicels (and leaves) minutely stipitate- 
glanduiar with gland-tipped hairs 0.1-0.3 mm, without villous hairs. Leaves mostly cauline (basal 

mostly deciduous by flowering), blades narrowly ovate or narrowly lanceolate to elliptic or elliptic- 



Nesom: Taxonomy of Erythranthe s 



lanceolate, largest 8-18(-30) mm x (l-)3-7 mm, relatively even-sized, or slightly reduced distally, 
palmately 3-5-veined, margins entire to mucronulate or denticulate, apex acute to obtuse, base 
attenuate to obtuse or rounded, narrowed to short (1-3 mm) petiolar regions on proximal, mostly 
subpetiolate to subsessiie distally. Fruiting calyces campanulate, maturing ovoid-ellipsoid to 
campanulate or broadly urceolate, 7-1 1 mm x 5-6 mm, winged and plicate-angled, distinctly inflated, 
sparsely and minutely hirtellous, eglandular, lobes subequal, 0.5-1.5 mm, ovate-deltate to broadly 
triangular, suberect, ciliate. Fruiting pedicels 7-18 mm, straight. Corollas yellow to pale yellow, 
sparsely red-spotted or not, tube-throats cylindric, 5-8 mm, exserted 1-2 mm beyond the calyx apex, 
limb weakly bilabiate, barely widened, lobes broadly obovate, apex rounded or mucronate. Styles 
glabrous. Anthers glabrous. Capsules oblong-ovoid to ovoid-cylindric or broadly ellipsoid, 5-9 
mm. Chromosome number unknown. Map 2. Afuller description is available in Meinke (1995a). 

Flowering Jun-Jul. Drying edges, banks, and beds of summer-dry watercourses, near drying 
edges of small lakes or impoundments, often among rocks and shoreline detritus, occasionally in 
moist protected areas beneath low shrubs (generally Artemisia tridentata), apparently restricted to the 
ecotone between the upslope edge of the sagebrush-juniper dominated shrub zone and the semi- 
aquatic graminoids near the water's edge (Meinke 1995a, b); 1200-1700 m. Calif, Idaho, Nev., 
Oreg. The type collection from Klickitat Co., Washingtoa was from a cultivated plant — no natural 
occurrences are known from Washington. 

Morphological and molecular data (Meinke 1995a; Beardsley et al. 2004) indicate that 
Erythranthe inflatula originated as a hybrid between E. breviflora and E latidens. If so, its 
geography and biology suggest that it is reproductively stable. The putative parents are 
geographically and ecologically saiarated for the most part and the range of E. inflatula is 
considerably broader than the relatively small region where the parents are sympatic. In this region, 
however, E. inflatula may be difficult to distinguish from one or both of its putative parents. 

Collections of Erythranthe inflatula are these, as cited by Meinke (2007) and with additions 
from the present stud}'. California. Lasse n Co. : 7.0 mi N of Madeline, ca. 1 mi N of Sage Hen siding 
on Southern Pacific RR, ca. 5500 ft, 28 Jun 1957, Bacigalupi 5989 (JEPS); 3.1 mi S of Madeline, 
ditches along US Hwy 395, on Madeline Plains, rich, black, peaty soil, 5300 ft, 28 Jun 1957, 
Bacigalupi 5998 (JEPS); 10 miles S of Ravendale, 9 Jun 1940, Pennell 25763 (P); 4,8 miles S of 
Madeline, 17 Jun 1958. Raven & Solbrig 13298 (JEPS). Modoc Co. : along Willow Creek, Jun 1894, 
Austin s.n. (UC); SW shore of Moll Reservoir, 2 meters from water's edge, 4920 ft, 5 Jul 1998, 
Schoenig 98-70 (DAV, JEPS); Damons Butte, ca. 2.6 mi W of Hwy 139, 4400 ft, cinder cone in a 
recent lava flow, in Pinus jeflreyi woodland with Artemisia tridentata understory, site recently 
burned, dominated by Ceanothus velutinus, 4 Jun 1988, Taylor 9745 (UC). ShastaCc^: ca. 1 mi W 
of Warner Grade Reservoir, on margins of N-most of four small seasonally inundated ponds, 3030 ft, 
14 Jun 1991, Taylor 1 1886 (UC). Sisky ou Co. : 0.8 mi E of Tennant Rd, NE of Weed, E of Grass 
Lake and W of Bray on Old State Hwy, ca. 1000 plants in scatteerd groups on a vernally wet, 
meadowy flat of low scrub, Oswald & Ahart 9359 (UC). Idaho. Owhyee Co. : meadow, 3 mi S of 
Riddle, 1 Jul 1949, Holmgren & Holmgren 7973 (CAS, UC, WS, WTU). Nevada. Washoe Co. : near 
CA-NV state line in S end of Coppersmith Hills, clay-loam vernal area with silver sage, 
Plagiobothrys, tansyleaf suncup, 6100 ft, 23 Jun 1986, Schoolcraft 1635 (UC). Oregon. Crook Co. : 
Grizzly Butte, 18 Jun 1894, Leiberg275 (NY, ORE, US). Gilliam Co. : forks of Cottonwood Canyon, 
6 Jun 1894, Leiberg 156 (NY, ORE, P, US). Grant Co. : Ochoco National Forest, Graylock Butte, 6 
Jul 1912, Ingram s.n. (RM). HarneyCa: dry watercourse near Frenchglen, 26 Jun 1942, Peck 21389 
(CAS, NY, P, UC, WILLI)) - probably on Burns BLM. Klamath Co. : along the E banks of Dog 
Hollow Reservoir, 12 Jun 2003, Meinke s.n. (OSC); shallow, stony drainage at SE edge of Campbell 
Reservoir, near culvert, Meinke s.n. (OSC) (both sites on Lakeview BLM). Lake Co.: along Dog 
Creek, W of Drews Reservoir, T. 40. S., R 17. E., Sec. 11 (NE %), 3 Jul 1999, Meinke s.n. (donated 
to Lakeview BLM herbarium; OSC); Sagehen Creek bed, just N of Road 4017 (west of Drews 



Nesom: Taxonomy of Erythranthe s 



Reservoir), T. 40. S, R. 17. E, Sec. 1 (SE %), 4 Jul 1999, Meinke s.n. (donated to Lakeview BLM 
herbarium; OSC); Whiskey Creek bed, above and below Road 4017 (just W of Drews Reservoir dam 
and picnic area), T. 40. S., R 18. E., Sec. 8 (NE %), 4 Jul 1999, Meinke s.n. (donated to Lakeview 
BLM herbarium; OSC); along Wool Lake drainage, SE margin of lake, along drying edges and banks 
of seasonal stream (mixed population with M. latidens), T. 38. S., R 25. E., Sec. 12 (NW %), 26 Jun 
1999, Meinke s.n. (donated to Lakeview BLM herbarium; OSC). Wasco Co. : Near Dalles City, 18 
Jun 1901, Suksdorfl029 (MO). Washington. The collection mapped from Klickitat Co. is the type. 

Meinke (2007) knew Erythranthe inflatula as represented by only five extant populations or 
population complexes — in Lassen Co., California, and Klamath and Lake cos., Oregon, but recent 
collections have substantiated its existence elsewhere and older localities should be revisited toward 
the possibility that the species persists there. The account of RTE species of Oregon (Oregon 
Biodiversity Information Center 2010) lists Malheur County in the distribution of the species. 
Oswald (1992) cited additional collections from Lassen Co., California: pool along base of RR, 
paralleling Poison Lake adjacent to Pittville Rd, Oswald 5652, 9263 (CHSC). 

8. Erythranthe latidens (A. Gray) Nesom, Phytoneuron 2012-39: 38. 2012. Mimulus latidens (A 
Gray) Greene, Man. Bot. San Francisco Bay, 278. 1891 Mimulus inconspicuus var. latidens 
A. Gray, Synopt. Fl. N. Amer. (ed. 2) (1, Suppl.): 450. 1886. LECTOTYPE: (cited by Grant 
1924, p. 202, as "TYPE"): USA, California. [Contra Costa Co.:] Near Monte Diablo, 26 May 
1862, W.H. Brewer 1161 (GH?; isotypes: UC!, US digital image!). The protologue gives "On 
the flanks of Monte Diablo, Brewer, Greene, and Chollas Valley, San Diego Co., Orcutt." 
The UC sheet is annotated as "Isotype!" by Grant. 

Grant (1924) indicated that she saw the Brewer collection at GH but it is not listed on 
the Harvard University Herbaria website. The specimen currently listed on the website as the 
type of Mimulus inconspicuus var. latidens is this: California. San Diego Co.: Chollas 
Valley, San Diego, 20 Jun 1884, C.R. Orcutt 679 (GH, MO!). An apparent duplicate of the 
Orcutt collection is at PH, although it has only incomplete collection data. On the Brewer 
1161 sheet at UC sheet, Pennell made the following annotation in 1941: "As type of M. 
inconspicuus latidens Gray I take C.R. Orcutt's 479, collected June 20, 1884 at Chollas 
Valley, San Diego, Calif., since it supplied Gray with most ample material and was his only 
collection acconip^mied by dissected flowers in a packet, thus showing its special study by 

Annuals, fibrous-rooted or filiform-taprooted. Stems ascending to ascending-erect, 3-10(- 
25) cm, usually multiply branched from the base. Stems arid pedicels (and leaves) short-stipitate- 
glandular to sessile-glandular with gland-tipped hairs 0.1-0.3 mm, without villous hairs. Leaves 
basal and cauline, largest at base or near midstem, sometimes unreduced in size up to the uppermost 
nodes, cauline blades ovate to ovate-lanceolate, 8-26(-35) mm, palmately 3(-5)-veined, margins 
entire or barely mucronulate to shallowly dentate-mucronulate, 1-3 teeth or mucronulae per side, 
apex acute to rounded, base abruptly cuneate to rounded, sometimes subauriculate, petioles absent. 
Flowers (l-)3-12. Fruiiting calyces tubular-campanulate, ovoid-ellipsoid, prominently 5-angled, 
purplish, 8-12 mm x 4-7 mm, strongly inflated, mostly minut^h stjpitak-^landul.k. lobes ^.ubajual 
triangular-acute, suberect, ciliate. Fruiting pedicels 11-28 mm. Corollas white to pinkish or flesh- 
colored, rarely yellowish, red-spotted on throat and lower lobes, tube-throats cylindric, 5-6(-8) mm, 
exserted 1-2 mm beyond calyx margin, limbs nearly actinomorphic, barely widened, lobes broadly 
obovate, rounded. Styles glabrous. Anthers glabrous. Capsules oblong to oblong-obovoid, 6-7 
mm. Chromosome number unknown. Map 2. 

Flowering Apr-Jun. Drained flats or slopes subject to vernal inundation, depressions in open 
fields, bare clay soil, vacant lots, roadsides; 10-800 m; Calif, Oreg.; Mexico (Baja California). 



Nesom: Taxonomy of Erythranthe s 



A collection of Erythranthe latidens from Madera Co., California, has fruiting calyces 8-12 
mm long and variably inflated. The distinction between E. latidens and E. inflatula sometimes seems 
arbitrary in northwestern California, where they are sympatric. 

9. Erythranthe moschata (Douglas ex Lindley) Nesom, Phytoneuron 2012-39: 38. 2012. Mimulus 
moschatus Douglas ex Lindley, Bot. Reg. 13: plate 1118. 1828 [1827?]. Mimulus guttatus 
var. moschatus (Douglas ex Lindley) Prov., Fl. Canada 1: 439. 1862. TYPE: USA. Oregon. 
[Multnomah Co.:] Garden specimens grown from seeds collected 'in moist rocks on the 
Multnomah and the Columbia at the falls of both, 23 May 1825, D. Douglas s.n. (holotype: 
BM?). A specimen at BM (Canada, 1826, D. Douglas s.n., BM digital image!) may prove to 
be the type but the collection data, as given by BM, do not appear to match the protologue. 
Lindley wrote that Douglas found it "growing sparingly on the margins of springs in the 
country about the river Columbia, in North-West America." 

Mimulus crinitus A.L. Grant, Ann. Missouri Bot. Gard. 11: 186. 1924. Mimulus acutidens Reiche, Fl. 
Chile 6: 63. 1911 (not M. acutidens Greene 1885). TYPE: CHILE. Prov. Valdivia, en 
pantanos, O. Buchtien 159 (holotype: SGO). 

Mimulus leibergii A.L. Grant, Ann. Missouri Bot. Gard. 11: 231, pi. 6, f 1. 1924. TYPE: USA. 
California. [Plumas Co.:] Mt. Pleasant, Spanish Peak Range, wet soil along creek, 6500 ft, 
16 Jul 1900, J.B. Leiberg 51 71 (holotype: US digital image! photo-MO! photo-UC!). 

Perennials, rhizomatous, rooting at lower nodes. Stems ascending to decumbent or 
prostrate, 2-30(^10) cm, simple or branched. Stems and pedicels glandular-villous with gland- 
tipped hairs 1-2 mm. Leaves cauline, basal not persistent, blades oblong-ovate, (10-)15^10(-50) 
mm x 5-25 mm, pinnately veined, margins coarsely serrate-dentate to denticulate or subentire, apex 
acute, base truncate to rounded or subcordate, petioles (l-)2-5(-10) mm. Fruiting calyces 
campanuiate, 6-13 mm, weakly inflated or not at all, villous to glandular-villous, plicate-angled. 
lobes lobes strongly unequal to subequal, 2-4 mm, triangular to linear-lanceolate or narrowly 
triangular-acuminate, erect to spreading-recurving, ciiiate. Fruiting pedicels (7-)10-25(-40) mm. 
Corollas mostly yellow with red to blackish or brown lines or red dots or both in the throat on the 
lobes, weakly bilabiate to nearly regular, tube-throats narrowly funnelform, 11-16 mm, lobes 
oblong-obovate, apex usually notched. Styles glabrous. Anthers glabrous to very slightly hirtellous 
or scabrous. Capsules ovoid, 3-7(10?) mm 2n = 32. Map 4. 

Flowering May-Aug. Shaded and wet places in sagebrush, aspen, spruce-fir, iodgepole pine. 
and meadows; 1200-3100 m; B.C., Alta., N.B., Nfdl., N.S., Ont, Que., P.E.I.; S.P.M.; [western 
USA] Calif., Colo,, Idaho, Mont., Nev, Oreg., Utah, Wash., Wyo., [eastern USA] Conn., Maine, 
Mass., Minn., N.H., N.J, NY, Penn, RI, Vt, Va, W.Va, Wis.; introduced: South America 
(Chile), Europe, Australia, New Zealand, Asia (Japan). 

Pennell (1935) noted that the Chilean Mimulus acutidens Reiche is the same species as the 
North American M. moschatus and indeed plants from Peru and Chile appear to be inseparable from 
M. moschatus in habit, vestiture, and calyx morphology. Penneil's assessment is corrobated in the 
present study, based on study of numerous South American collections at MO and LIT. Fruiting 
calyces of the Chilean plants are 6-8 mm, barely expanded from flower; corolla tube-throats 10-12 
mm. Von Bohlen (1995b) maintained Mimulus crinitus as a distinct species (including M. acutidens 
Reiche as a synonym) but noted that a closer analysis of North American material of M. moschatus 
would be necessary for a better judgement. Mimulus acutidens and M. moschatus sensu stricto are 
conspecific. 

Erythranthe moschata as treated here, recognizing the segregates E. moniliformis and E. 
inodora, is narrower in concept than those of recent decades (e.g., Thompson 1993; Carlson 2002; 
Whittall et al, 2006), where only the single broad species was recognized. Whittall et al. (2006) noted 



Nesom: Taxonomy of Erythranthe s 



that "the Mimulus moschatus alliance is a group of 13 closely related species with uncertain species 
boundaries and interspecific relationships (Grant 1924; Pennell 1951; Argue 1986; Meinke 1992; 
Whittall 1999; Carlson 2002)," but this seems unfair particularly to Pennell, who recognized the two 
segregates of M. moschatus (he also recognized M. macranthus as distinct, but it is here regarded as 
conspecific withM moniliformis). Studies of pollen morphology (Argue 1980, 1986) also explicitly 
support the segregation ofM moniliformis (which has unormamented muri) fromM moschatus. 
Argue's tentative distinction (1980) of the pollen of M. inodorus from that of M. moschatus was not 
confirmed in the 1986 study. 

In contrast, it seems that neither Meinke nor Whittall nor Carlson lias even mentioned the 

potential existence of the segregates. Munz and Keck (1959) recognized Mimulus moschatus var. 
moniliformis as distinct but treated M. inodorus as a synonym of M. moschatus var, moschatus. The 
only synonym included by Thompson (1993) as a synonym of the broadly conceived M. moschatus 
was "M. moschatus var. moniliformis." 

Each of the three species recognized here (Erythranthe moschata, E. moniliformis, E. 
inodora, using criteria similar to those of Pennell) has a distinct range but each overlaps with the 
other two (Maps 3 and 4), and while there are indications that hybrids may be formed, all three 
apparently maintain their morphological distinction. Zones of intergradation in the areas of overlap 
are not evident but field studies m~q needed to examine this more closely. 

10. Erythranthe moniliformis (Greene) Nesom, Phytoneuron 2012-39: 38. 2012. Mimulus 
moniliformis Greene, Bull. Calif. Acad. Sci. 1: 10. 1884. Mimulus moschatus var. 
moniliformis (Greene) Munz, Aliso 4: 99. 1958. TYPE: USA. California. [Kern Co.:] 
Summit Station, Aug 1883, E.L. Greene s.n. (holotype: CAS?; isotypes: GH, MO!). Not 
located at ND-Greene. Protologue: "Common in dry rocky places of the Sierra, from 4,000 to 
8,000 ft. In the collections of Bolander, Kellogg and others, this species occurs abundantly, 
and is named M. moschatus,' being confounded with the last species." 

Mimulus dentatus var. gracilis A Gray, Bot. Gaz. 7: 112. 1882. TYPE: USA. California. [Shasta 
Co.:] Lassen's Peak, 1882, Mrs. R.M. Austin s.n. (holotype: GH). Not located at ND-Greene. 

Mimulus moschatus var. sessilifolius A Gray, Synopt. Fl. N. Amer. (ed. 2) 2(1): 447. 1886. Gray- 
noted as a synonym "M. inodorus Greene, I.e., but the plant as strongly musk-scented as the 
ordinary species, at least in some cases." Gray also noted "Not rare in wet places, from San 
Bernardino Co., California, northward, and passing into the ordinary form in Oregon." 
SYNTYPES: USA. California: Butte Co.: Chico, Apr 1885, A. Gray s.n. (GH). California. 
Alameda Co.: Temescal, Tule swamps, Jun 1885, W.S. Lyon <5(GH). 

Mimulus moschatus var. longiflorus A Gray, Synopt. Fl. N. Amer. (ed. 2) 2(1): 278, 1886. Gray- 
noted only "The usual form in California, also in Oregon." In a later citation (p. 447), he 
noted "M. moniliformis, in part (the villous- and more or less viscous-pubescent plant), 
Greene, Bull. Calif. Acad. i. 119. Common especially in the Sierra Nevada." SYNTYPES: 
USA. California. Mariposa Co.: Yosemite Valley, 1886, H.N. Bolander 6306 and 6307 
(GH). 

Mimulus macranthus Pennell, Proc. Acad. Philad. 99: 160. 1947 TYPE: USA. California. Shasta 
Co. : Hatchet Mountain, 6 to 8 mi W of Burney, along stream in coniferous (Pseudotsugd) 
forest, 3900 ft, 7 Jun 1940, F. W. Pennell 25710 (holotype: PH!). 

Perennials, rhizomatous. Stems erect to ascending, 10-30 cm, simple or usually branched. 
Stems and pedicels (and leaves) densely villous with eglandular or weakly gland-tipped hairs 0.5-2 
mm, very rarely glabrate. Leaves mostly cauline, basal not persistent, blades oblong-ovate to ovate, 
18^0 mm x 7-13 mm, pinnately veined, margins dentate to denticulate, apex acute to obtuse, base 
rounded to subcordate, subclasping to sessile, petioles absent or 0.5-1 mm. Fruiting calyces 



Nesom: Taxonomy of Erythranthe s 



cylindric-campanuate, 10—1 1(— 13) mm, weakly inflated, ridged-angled to winged-angled, vilious- 
glanduiar, lobes subequal, 2-3 mm, lanceolate to triangular-subulate, spreading-recurving, ciliate. 
Fruiting pedicels 10^10 mm. Corollas yellow with fine blackish or brownish lines on all sides of 
the throat, red to brown spots present or not, tube-throats cylindric-funnelform, 12-18 mm, limbs 
weakly bilabiate to nearly regular, apex rounded. Styles glabrous. Anthers glabrous, rarely 
hirtellous. Capsules narrowly elliptic-ovoid, 6-8 mm. Chromosome number unknown. Map 4. 

Flowering Jun-Aug. Around springs and seeps, creek edges, moist meadows, ditches, along 
trails, roadsides, rocky ridges, granite outcrops, serpentine talus, fir and pine forests; (1000-)1500- 
2800 ft; Calif., Oreg. 

In his description of Mimulus macrcmthus, Pennell (1947, p. 160-161) noted that "This 
comprises the major part of the material that has been called Mimulus moschatus longiflorus Gray 
(not M. longiflorus (Nutt.) Grant), the remainder being mostly M. inodorus Greene. These plants, 
especially developed in the Cascade Range, and includingM moniliformis Greene, have been usually 
treated as forms of the Musk Flower, M. moschatus Dougl., but they seem better considered as 
distinct species ... ." He recognized both M. macranthus and M. moniliformis, distinguishing the 
latter by its "finely pubescent to glabrous leaves" and petiolate leaves. The vestiture proves to be 
more variable than allowed in Pennell's concept, thus Greene's older epithet has priority for this Sierra 
Nevada-centered species. 

Mimulus moniliformis was described by Greene as "Near M. moschatus, wholly scentless, 
villous but scarcely viscid, 3-8 inches high from a perennial root, with subterranean shoots bearing 
moniliform strings of small tubers." Production of odor has been noted by various collectors and 
authors to vary in Erythranthe moschata and slender "moniliform" rhizomes also are variably 
produced within the species — neither feature separates E. moniliformis from E. moschata. 

Erythranthe moniliformis is distinct from E. moschata in its erect habit (vs. decumbent to 
procumbent in E. moschata) and characteristically sessile to subsessile cauline leaves (vs. usually 
petiolate in E. moschata), lending an easily recognizable aspect to the plants. Pedicels of E. 
moniliformis are ascending-erect, while in E. inodora and E. moschata, both of which are essentially 
prostrate, pedicels are characteristically spreading at about 90 degrees. Leaves of E. moniliformis 
sometimes are short-petiolate and the distinctions in vestiture and corolla size noted by Pennell 
(1947) are not consistent. In spite of what may appear to be subtle differences, the two are distinct in 
geography. 

11. Erythranthe inodora (Greene) Nesom, Phytoneuron 2012-39: 38. 2012. Mimulus inodorus 
Greene, Bull. Calif. Acad. Sci. 1: 119. 1885. Greene did not cite a type; he referred to "M. 
moschatus, Gray, Bot. Cal. I. 569, not of Dougl. M. moschatus, var. longiflorus, Gray, Syn. 
Fl. 278" and noted "Common in both the Coast Range and the Sierra Nevada, throughout 
California, and also in Oregon." In the Synoptical Flora, Gray noted only "The usual form in 
California, also in Oregon." 

Grant (1924) recognized this entity as Mimulus moschatus var. sessilifolius A Gray 
(with M. inodorus as a synonym), here placed as a synonym of Erythranthe moniliformis, and 
noted that it occurs from British Columbia to southern California. She cited numerous 
specimens but not a type. Pennell (1951) treated it at specific rank, describing its range in 
California as only in northern counties. 

Mimulus moschatus var. pallidiflorus Suksdorf, Deutsche Bot. Monatsschr. 18: 154. 1900. TYPE: 
USA. Washington. Skamania Co.: Springs near Chenowith, 7 Jul 1894, W.N. Suksdorf 2320 
(holotype: ?; isotypes: ORE digital image!, UC, US digital image!). 



Nesom: Taxonomy of Erythranthe s 



Perennials, rhizomatous, sometimes rooting at lower nodes. Stems usually prostrate, 
sometimes decumbent to ascending, 20-80 cm, few-branched. Stems and pedicels (and leaves) 
villous with eglandular hairs 1-2 mm, sometimes mixed with much shorter stipitate-giandular hairs. 
Leaves cauline, basal not persistent, blades oblong-lanceolate, 30-70 mm x 10-22 mm, pinnately 
veined, margins denticulate to dentate, apex acute, base rounded, petioles absent or uncommonly with 
petioles l-2(-3) mm. Fruiting calyces cylindric-campanulate, 10-12 mm, weakly inflated, wing- 
angled or plicate-angled, glandular-villous with gland-tipped hairs, lobes strongly unequal, 5-9 mm, 
linear-lanceolate to narrowly triangular with long-acuminate-apiculate apices, spreading, ciliate. 
Fruiting pedicels (15-)22-50 mm. Corollas yellow with line blackish or brownish lines on all sides 
of the throat, tube-throats narrowly campanulate, 15-18 mm, limbs weakly bilabiate and often nearly 
regular, apex rounded. Styles glabrous. Anthers consistently finely hirtellous to hispidulous. 
Capsules 6-8 mm. Chromosome number unknown. Map 5. 

Flowering Jun-Sep. Creek banks, gravel bars, flood plains, shallow ditches, swales, damp 
banks, moist soil in coniferous woods, marshes, bogs, wet sand; 0-1900 m; British Columbia; Calif, 
Oreg., Wash. 

Greene (1885) characterized the species as "Quite distinct from the true musk plant, being of 
more than twice the size, scentless, and possibly only animal; certainly never rooting at the joints," 
further noting "villous and slimy but wholly scentless; stems 1-3 feet long, weak and decumbent, but 
not creeping or rooting; leaves ... closely sessile by a broad base." In contrast to Greene's 
observation, several collections examined in the present study show that it indeed may root 
adventitiously at lower nodes. 

Erythranthe inodora is recognized by its prostrate to decumbent or decumbent-ascending 

habit, large, mostly sessile leaves, dense villous vestiture, long pedicels, large calyces and corollas, 
hispid-hirtellous anthers, and particularly by its very long, strongly unequal, linear-triangular calyx 
lobes. The leaves typically and characteristically are sessile with truncate to rounded or subcordate 
bases but some are short-petiolate (e.g.: California: Lake Co., Heller 5923: Siskyou Co., Heller 7960 
and Oettinger 478. Oregon: Gilliam Co., Jones 28831; Jackson Co., Hammond 312. Washington: 
Skamania Co., the type of Mimulus moschatus var. pallidiflorus). In these cases, the distinctive leaf 
bases, vestiture, calyx morphology, and pubescent anthers are generally diagnostic. 

A population system of Erythranthe inodora-like plants from counties in southern California 
probably was the basis for Pennell's attribution (1951) of the species to that area, apparently about 
300 miles disjunct from the main range of the species. These plants have the prostrate habit, large 
leaves, long pedicels, and large corollas of E. inodora but the calyx lobes are variable in length and 
usually do not show the attentuate-apiculate apices characteristic of the latter. Representative 
specimens: California. Los Angeles Co. : Verdugo Canyon, damp woods, 11 Aug 1910, Blake 1639 
(LL); Oak Knoll, Aug 1902, Braunton 656 (UC). San Bernardino Co. , San Bernadino Mts.: marsh, 
4000 ft, 21 Jul 1897, Chandler s.n. (UC); Lower South Fork Meadow, San Gorgonia Wilderness 
Area, grassy montane meadow r in mixed conifer forest, mostly Pinus Jeffrey i andAbies concolor, ca. 
7600-8000 ft, 15 Aug 1976, Davidson 4713 (UC); 1.2 mi E of Running Springs (town) on Calif. 18, 
common along seepage in road cut, sandy soil, 6100 ft, 19 Jun 1969. Holmgren 3596 (UC); Bluff 
Lake, shade of willows. 7400 ft, 13 Jufl926, Mum 10690 (UC); Little Bear Valley, Aug 1884, 
Parish 1463 (UC-2 sheets); 1 mi ESE of Jenks Lake, Santa Ana R. drainage, shady creek bed, 7000 
ft, 20 Aug 1932, Wheeler 1158 (UC). San Diego Co. : Palomar Ml, 5500 ft, 7 Apr 1928, Meyer 504 
(UC); Palomar Ml, near Palomar Hotel, 4 Jul 1928, Meyer 504 (JEPS). Riverside Co.: San Jacinto 
Mts, Tauquitz Valley, wet meadow, 7500 ft, 8 Aug 1903, Jepson 2300 (JEPS); Hannah's Sawmill, 
1/2 mi E of Dutch Flat, edge of stream, ca. 6000 ft, 26 Jul 1928, Meyer 524 (UC 2 sheets); San 
Jacinto Mts, Tahquitz Valley, on a stream bank, 3 Jul 1928, Meyer 601 (JEPS, UC). 



Nesom: Taxonomy of Erythranthe s 



Plants seemingly disjunct inland in Butte Co., California, are otherwise typical of the species. 
Examples : Butte Co . : Damp sand at S edge of Butte Creek, ca. 2. 1 mi S of Skyway and ca. 1/4 mi E 
of Hwy 99 bridge, 29 Aug 1987, Castro 213 (DAV); Chico, 28 Jul 1916, Hazeltine s.n. (DAV). 

12. Erythranthe pulsiferae (A. Gray) Nesom, Phytoneuron 2012-39: 38, 2012. Mimulus pulsiferae 

A. Gray, Proc. Amer. Acad. Arts 11: 98. 1876. LECTOTYPE (Grant 1924, p. 212): USA. 
California. [Plumas Co.:] Indian Valley, 1873, Mrs. Pulsifer-Ames 21 (GH). Gray cited 
"California, in the Sierra and Indian Valleys of the Sierra Nevada, Bolander, Mrs. Pulsifer- 
Ames." Grant (1924) cited Ames 21 as the "Type." 

Annuals, shallow!}' fibrous-rooted. Stems erect, 5-12(-18) cm, simple or sparingly 
branched at the base. Stems and pedicels minutely stipitate-glandular with gland-tipped hairs 0.1- 
0.3 mm. Leaves basal and cauline, blades elliptic-oblong to ovate or oblanceolate, 3-14(-23) mm x 
2-9(-15) mm, palmately 3-veined, margins denticulate to entire, apex acute to obtuse, base cuneate to 
attenuate, petioles 2-9 mm. Flowers 1-5. Fruiting calyces cylindric, 7-10 mm, weakly to strongly 
inflated, stipitate-glandular, lobes subequal, 0.9-1.1 mm, triangular-acute, suberect, ciliate. Fruiting 
pedicels 12-38 mm, divergent-arcuate. Corollas yellowish, "limb pale yellow with pink edges, 
throat yellow," "white with yellow throat and pink border to the expanded limb," reddish dots present 
or not on the lower lip, tube-throats funnelform, 6-9 mm, limbs weakly bilabiate, lobes broadly 
obovate-suborbicuiar, apex rounded. Styles glabrous. Anthers glabrous. Capsules fusiform- 
cylindric, 5-8 mm. In = 32. Map 6. 

Flowering Apr-Jul. Damp depressions, moist gravel, rocky flats, granite outcrops, wet 
meadows, lava beds, vernal pools, forest openings, commonly in or near coniferous forest, also 
chaparral-live oak; 50-1300(-2500) m; Calif., Oreg., Wash. 

Erythranthe pulsiferae is characterized by its minutely stipitate-glandular vestiture (lacking 
villous hairs), elongate internodes, persistent basal leaves, small, palmately veined, short-petiolate 
cauline leaves with elliptic-oblong to ovate or oblanceolate blades, divergent-arcuate pedicels, and 
small, weakly bilabiate corollas. Erythranthe floribunda is distinct from E. pulsiferae in its 
multicellular vestiture, urceolate fruiting calyces, and pinnately veined leaves with generally deltate 
to ovate blades and more strongly toothed leaf margins. The vestiture of E. pulsiferae is generally 
more similar to that of the Columbia River clade. 

The anther pairs and stigma of Erythranthe pulsiferae are at essentially the same level and the 
species appears to be consistently autogamous over its range. Rare plants in Humboldt County (e.g., 
Tracy 761 6 and 12S38, JEPS) have slightly longer corolla tube-throats and broader limbs — in these 
the anther pairs are slightly separated and the stigma is at or slightly above the upper anther pair. 

13. Erythranthe arenaria (AL. Grant) Nesom, Phytoneuron 2012-39: 38. 2012. Mimulus arenarius 

AL. Grant, Ann. Missouri Bot. Gard. 11: 215. 1924 [1925]. TYPE: USA. California. Fresno 

Co.: moist sandy places near Huntingdon Lake, 7000 ft, 5 Jul 1917, A.L Grant 1032 

(holotype: MO!; isotypes: DS, GH, BH, JEPS!, OS digital image!, PH!, POM, RM US 

digital image!). 
Mimulus subulatus (AL. Grant) Pennell, Proc. Acad. Nat. Sci. Philad. 99: 162. 1947. Mimulus 

floribundus var. subulatus AL. Grant, Am. Missouri Bot. Gard. 11: 222. 1924. TYPE: USA. 

California. [Tuolumne Co.:] between Hog Ranch and Hetch-Hetchy Valley, 4200 ft, 16 Jun 

1917, A.L Granf 970 (holotype MO!; isotypes: GH, JEPS!, US digital image!). 
Mimulus multiform Pennell, Proc. Acad. Nat. Sci. Philad. 99: 161. 1947. TYPE: USA. California. 

Fresno Co.: 4 mi E of Dunlop, moist granitic gravelly sand, 3700-3800 ft, 9 Aug 1940, F.W. 

Pennell 26451 (holotype: PH!; isotypes: MO!, NY 2 sheets digital images!, US digital 

image!). 



Nesom: Taxonomy of Erythranthe s 



Mimulus trisulcatus Pennell, Hoc. Acad. Nat. Sci. Philad. 99: 161. 1947. TYPE: USA. California. 

[Tulare Co.:] below Mineral King [western slope], moist gravelly granitic soil, 7000-7600 ft, 

6 Aug 1940, F.W. Pennell 26421 with A. Cronquist (holotype: PH!; isotypes: MO!, NY 2 

sheets digital images!, UC!, US digital image!). 

Annuals, fibrous-rooted or filiform-taprooted. Stems erect to ascending, 5-20 cm, simple or 
branched. Stems and pedicels villous-glandular with gland-tipped hairs 0.2-0.8 mm. Leaves basal 
and cauline, blades elliptic to narrowly elliptic, ovate-elliptic, or ovate-lanceolate, 5-12(-17) mm x 
3-7 mm, 1-veined or palmately 3-veined, margins entire to sparsely dentate to serrate, apex 
acuminate to acute or obtuse, base rounded to cuneate-attenuate, petioles essentially absent or 
proximally l-3(-5) mm on proximal leaves. Fruiting calyces narrowh ..ainpanulate 5-7(-9) mm, 
not inflated or weakly so, often red-dotted, villous-glandular, lobes subequal, ca 1 mm, deltate- 
subulate to broadly triangular, suberect ciliate. Fruiting pedicels 10-23 mm, divergent-arcuate. 
Corollas yellow with red-mottled lower lip, tube-throats funnelform, 9-12(-14) mm, limbs weakly 
bilabiate, lobes broadly obovate, apex rounded. Styles glabrous. Anthers glabrous. Capsules 
elliptic, 4-7 mm In = 32 (Heckard 4067, JEPS). Map 6. 

Flowering May-Sep. Sandy flats, bars, gullies, washes, trails, and road cuts, seasonal creek 
beds and drainages, rocky slopes, seepy loam, ditches, lake edges, meadows, openings in pine-fir and 
pine-oak woodlands; (100-)500-2600(-2800) m; Calif. (Fresno, Los Angeles, Madera, Mariposa, 
Tulare, Tulomne). 

Most plants of Erythranthe arenaria have relatively even-sized cauline leaves, all sessile to 
subsessile. Some, however, have persistent basal leaves that are short-petiolate, ovate with a cuneate 
base, and relatively larger than the more distal cauline ones. Such plants are those named by Grant as 
Mimulus floribundus var. subulatus. These might be construed as showing influence of E. 
geniculata, but the latter occurs only at the lower range of elevation for E. arenaria and the 
"subulata" variants occur at least up to 2300 meters. The "subufafa" variants also have the erect habit 
characteristic of E. arenaria. In any case, these variants should be investigated, especially in the 
Yosemite area where they appear to be relatively common, toward the possibility that they represent a 
distinct entity. 

It is remarkable that Pennell described two species, based on his own collections and field 
observations, that appear to be essentially segregates from Erythranthe arenaria. Yet there do not 
appear to be discontinuities in corolla size or morphology, features he emphasized in descriptions of 

Mimulus multiflorus and Mimulus trisulcatus. Variability in overall size perhaps contributed toward 
recognition of M. multiflorus. California botanists perhaps will be able to corroborate Pennell's field 
observations of corolla variation and partition the variation more precisely. 

Plants of a collection of Erythranthe arenaria from Mariposa County (N of Fish Camp, 4900- 
5100 m, Pennell 26392, UC) have mature calyces conspicuously longer (8-9 mm vs 5-7 mm) and 
more inflated than characteristic for the species and the corollas are slightly longer. Otherwise, they 
seem securely identified as E. arenaria. 

Plants of the apparently disjunct collection from Los Angeles County are similar to those in 
counties further north: Los A ngeles Co.: San Gabriel Mts., upper slopes of Little Rock Creek Canyon, 
ca. 1 mi W of Cedar Spring, sandy base of scree slope on N face of Kratke Ridge, 6800 ft, 20 Jul 
1958, Bacigalupi 6416 (JEPS). 

14. Erythranthe floribunda (Douglas ex Lindley) Nesom, Phytoneuron 2012-39: 38. 2012. 
Mimulus floribundus Douglas ex Lindley, Bot. Reg. 13: plate 1125. 1827. TYPE: USA. 
Washington. Protologue: "A neat hardy annual, found by Mr. Douglas on moist rocks in the 
interior of the districts of the river Columbia" (holotype: K?), Grant (p. 218) noted "on 



Nesom: Taxonomy of Erythranthe s 



limestone rocks on dry sandy soils in the interior of the Columbia, 1826, Douglas (G, 
probably part of the type collection)." 

Mimulus peduncularis Douglas ex Bentham, Scroph. Ind., 29. 1835. TYPE: USA. "America boreali- 
occident," 1826, Douglas s.n. (holotype: K, sketch and fragment of type at MO!). The full 
entry of the protologue is this "21. M. PEDUNCULARIS (Dougl. MSS.), pubescens, humilis, 
foliis petiolaris ovatis acutis subdentatis basi cuneatis rotundatisve, calycibus (parvis) ovato- 
tubulosis, dentibus brevibus acutis subqequalibus. — America boreal i-occident. Douglas." 
Collected in 1826, D. Douglas s.n, (holotype: K, sketch of type at MO). 

Mimulus deltoideus Gandoger, Bull. Soc. Bot. France 66: 218. 1919. TYPE: USA. Oregon. [Lake 
Co.:] North Pine Creek, near Snake River, moist situations, 13 Jul 1899, W.C. Cusick 2237 
(holotype: ?; isotypes: BH, MO!. OS digital image!, UC!). 

Mimulus serotinus Suksdorf, Deutsche Bot. Monatsschr. 18:154. 1900, TYPE: USA. Washington. 
Klickitat Co.: damp sandy banks of the Columbia River, Oct-Dec 1892, W.N. Suksdorf 2185 
(holotype: WS; isotypes: DS, MO!, NY digital image!, ORE digital image!. UC 2 sheets!, US 
digital image!; possible isotype: NY digital image!). 

Mimulus membranaceus A. Nelson, Bot. Gaz. 34: 30. 1902. Mimulus floribundus var. membranaceus 
(A. Nelson) A. L. Grant, Ann. Mssouri Bot. Gard. 11: 221. 1924. TYPE: USA. Wyoming. 
[Albany Co.:] Centennial Hills, 16 Jul 1894, A. Nelson 1683 (holotype: RM; isotypes: BH, 
GH. MO!, NY digital image!). 

Annuals, fibrous-rooted or filiform-taprooted. Stems 3-22(-40) cm, erect to decumbent, 
sometimes procumbent-trailing, simple to many-branched. Stems and pedicels villous-glandular 
with gland-tipped hairs greatly variable in length and density, sometimes reduced to sparsely 
stipitate-glandular with hairs 0.2-0.5 mm. Leaves cauline, basal mostly deciduous by flowering, 
blades ovate, (3-)8-25(-35) mm x ( 1 - ) 5 — 1 8 (—26 ) mm, pinnately to subpalmately veined, margins 
serrate to sparsely dentate, apex acute, base cuneate to truncate or cordate, petioles 1-12 mm. 
Fruiting calyces cylindric, 4-7 mm, weakly to strongly inflated, greenish or purplish to red-dotted, 
villous-glandular, lobes subequal, (0.5-)0.8-1.6(-2) mm, triangular-acuminate, suberect, ciliate. 
Fruiting pedicels 5-20(-26) mm. Corollas yellow with red-dotted lower lip, tube-throats 
funnelform-cylindric, (4-J5-10 mm, limbs weakly bilabiate, expanded 3-4 mm across (pressed), 
lobes mostly oblong, apically notched. Styles glabrous. Anthers glabrous. Capsules obovoid to 
elliptic, 3.5-7 mm. 2n = 32. Map 7. 

Flowering (Apr- in Arizona, May-)Jun-Aug(-Sep). Under overhangs, moist roofs of cave 
ruins, w ? et rock crevices, cliff faces and wet cliff bases, below waterfalls, seeps, springs, humus and 
moist soil over rocks and slabs, moist slopes, along ditches and pond edges, wet edges of creeks and 
rivers, drying mud on margins of wetland depressions, creek beds, wet or swampy meadows, along 
trails, in lodgepole pine, ponderosa pine, ponderosa pine-douglas fir, and spruce-fir woodlands; 
(1000-)1800-1600(-3100) m (ca. 300-500 m in Arkansas); Alta., B.C.; Ariz., Ark, Calif., Colo., 
Idaho, Mont, Nev., N.Mex., Oreg., Wash., Utah, S.Dak., Wyo.; Mexico (Baja California, Baja 
California Sur, Chihuahua Smaloa. Sonora). 

Collections have been made of plants much reduced in size — in leaves, flowers, and overall 
stature — so strikingly so that one might suspect that they are evolutionarily distinct, but the sizes 
appear to be at the lower limits of the species (as in the description above) and such plants are 
identified here as Erythranthe floribunda. The following is an example: Nevada. Nye Co. : Toquima 
Range, Toiyabe National Forest, Pine Creek Canyon, ca. 8500 ft, abundant along small stream, 13 Jul 
1964, Holmgren and Reveal 1444 (TEX). 

Erythranthe floribunda has been recognized from northern Arkansas (e.g., Moore 1958), 
where documented from a number of counties (Carroll, Cleburne, Crawford, Franklin, Izzard, 
Johnson, Logan, Newton, Pope, Searcy, Stone, and Washington). The unpublished name "Mimulus 
floribundus subsp. moorei litis" has appeared in various checklists in reference to the Arkansas plants, 



n: Taxonomy of Erythranthese 



but observations in the current study of populations in the herbarium and field indicate that they are 
not distinct from the rest of the species. The disjunction in geography appears to be analogous to that 
in El moschata. 



Some plants identified here as Erythranthe floribunda in Arizona and southwestern New 
Mexico (e.g., Figs. 1--3) are distinctive in their prominently inflated calyces, sessile to subsessile 
leaves with attenuate bases and palmately 3-5-nerved venation, and much- elongated pedicels (20— 43 
mm), but intermediates in Arizona make it difficult to conclude that the variants represent an entity 
discontinous from plants of typical morphology. This variant morphology has not been observed 
in .--i.j i. i. .I. |--|-ul ■■■- -ii 




Figure 1 . Erythranthe floribunda 
Worthington 32511, SRSC). 



i: Taxonomy of Erythranthe st 




Figure 2. Erythranthe ft oribunda variant from Hidalgo Co., New Mexico (Todsen s.n., NMC). 



i: Taxonomy of Erythranthe st 




Figure 3. Erythranthe floribunda variant from Williams, Coconino Co. (Greene s.n., ND-Greene). 



Nesom: Taxonomy of Erythranthe s 



1 . Cauline leaves basally attenuate to sessile or subsessile, epetiolate, blades oblanceolate to elliptic or elliptic- 
lanceolate, primarily palmately 3-5-nerved, sometimes with an additional 1-2 smaller lateral pairs; fruiting 
pedicels 20-43 mm; corolla tube-throats 5-6 mm; fruiting calyces 5-8 mm 

Arizona/New Mexico variant 

1. Catiline leaves abruptly and distinctly petiolate, blades generally ovate with a rounded to truncate or cordate 
base; mostly pinnately to subpalmately veined; fruiting pedicels 5-15(-26) mm; corolla tube-throats (4-)5-10 
mm, fruiting calyces 4-7 mm Erythranthe floribunda 

Collections examined of the Erythranthe floribunda variant. Arizona. Coconino Co. : 
Williams, wet meadow, 6 Jul 1889, Greene s.n. (ND-Greene 2 sheets). Pima Co. : S end of 
Baboquivari Mts., in sand along edge of Presumido Wash near Buenos Aires, ca. 3000 ft, 3 Apr 1966, 
Mason 2559 (ARIZ). Santa Cruz Co. : Meadow Hills Country Club, 4.5 mi N of Nogales, wet soil in 
marsh, [ca. 3900 ft,] 5 May 1966, Crutchfield 1460 (XL); Santa Rita Mts,, 4500 ft, 17 Apr 1903, 
Thornber 505 (UC). New Mexico. Hidalgo Co. : 30 mi S of Animas, edges of ponds near Gray's 
Ranch and sandy banks of Animas Creek, 5000 ft, 19 Jun 1973, Todsen s.n. (NMC). Peloncillo Mts., 
Clanton Draw, 3.0 mi E of the [Coronado] National Forest, E side property line, 5480 ft, annual at 
edge of stream, 9 May 2004, 111). Worthington 3251 1 (NMC, SRSC, UNM, UTEP). 

Previous tentative identifications on the three Hidalgo County sheets include Mimulus 
floribundus, M. "probably floribundus," M. primuloides, and M. rubellus. Similar ambiguity in 
identification of New Mexico collections recently confirmed as Erythranthe suksdorfii was noted by 
Keller (2010), but the New Mexico distribution of this species is north of Hidalgo County and the 
collections cited by Keller seem securely identified as E. suksdorfii (fide Phil Tonne, UNM). 

A collection from southwestern Mexico is similar to Erythranthe floribunda but it is far- 
disjunct from other populations of that species, completely prostrate, and has very short pedicels: 
Jalisco. W of San Sebastian, Hacienda del Ototal, wet sand of stream bottom, 1500 m, Alexia 1853 
(UC). Fig. 4. It perhaps belongs with plants named by Bentiiam as Mimulus pubescens, which may 
prove to be a distinct species. 

Mimulus pubescens Bentham, Prodr. (DC.) 10: 372. 1835. TYPE: MEXICO. Jalisco. "In Mexico 
prope Talisco," Beechey s.n. (holotype: K, photo MO!). The only information on the 
specimen is "Mexico, Beechey." See comments below. 



r. Taxonomy of Erythranthes 




Nesom: Taxonomy of Erythranthes 



15, Erythranthe austrolatidens Nesom, sp< nov. TYPE: MEXICO. Baja California Sur. Sierra 

Guadalupe, W of Mulege, W side of the mountain range, vicinity of Rancho El Tule, E of San 
Martin and La Vinorama, 26° 8 1' N, 1 12° 72' W, rocky volcanic substrate, with Lysiloma 
Candida, Opuntia cholla, Pachycereus pringlei, ca. 260 m, 26 Apr 1998, Rebman 51 70 (SD!). 

Similar to Erythranthe latidens in its annual duration, fibrous roots, yellow, cleistogamous/ 
autogamous flowers, sessile leaves, tubular campanulate calyces with 5 equal, deltate lobes: different 
in its short villous -glandular vestiture of stems, leaves, pedicels, and calyces, petiolate proximal 
leaves, consistently serrulate-denticulate leaf margins, generally smaller fruiting calyces, and disjunct 
geography in Baja California Sur. 

Additional collections examined. MEXICO. Baja California Sur. Mpio. Comondu, La 
Laguna, al norte de San Jose de Comondu, matorral sarcocaule, 443 m, 26° 0& 48.2" N, 111° 46' 
38.2" W, 14 Mar 2002, DominguezL. 3136 (ARIZ-2 sheets, SD). Mpio. Comondu, Sierra La 
Giganta, Llanos de San Julio, 5 km al E de San Jose de Comondu, matorral xerofilo, 432 m, 19 Feb 
2003, Dominguez L. 3399 (.ARIZ). Mpio. de La Paz, Mison de Los Dolores, 3 km al S W del Rancho 
Los Dolores, 25° 03' 20.8" N, 110° 53' 28.1" W, matorral xerofilo, 85 m, 15 Mar 2003, Dominguez L. 
3448 (SD). 

Annuals, fibrous -rooted. Stems terete, 6-22 cm, nodes 3-5, branched mostly at the base, 
erect to erect-ascending. Stems, leaves, pedicels, and calyces short-villous- glandular with gland- 
tipped hairs 0. 1-1.0 mm, without egiandular hairs. Leaves basal and cauline, basal often deciduous 
by flowering, largest at base or near midstem, cauline blades ovate to elliptic or obovate, 5-25 x 2-11 
mm, basal blades to 30 mm long, 15 mm wide, palmately 3-6-veined, often suprabasal, margins 
consistently s errata te-mucronulate to -denticulate, (2-)4-6 teeth per side, apex sharply acute, base 
acute to cuneate, basal and lower cauline petiolate with petioles 1-6 mm. Flowers 5-13, from basal 
to distal nodes. Fruiting calyces tubular-camp anulate, ovoid-ellipsoid, 5 angled, greenish with 
purple ribs, 8-9 mm x 3-5 mm, slightly inflated, lobes subequal, deltate-rounded, apex apiculate, 
margins weakly ciliate. Fruiting pedicels 5-30 mm. Corollas yellow, apparently without red 
markings, tube-throats cylindric, 7-8 mm, exserted 2(-3) mm beyond calyx margin, limbs nearly 
actinomorphic, barely widened, lobes obovate. apices rounded. Styles glabrous. Anthers glabrous. 
Capsules 4-6 mm, oblong to oblong-ob ovoid, stipitate. Map 3. 

Flowering (Jan-)Feb-Apr. Xeric shrubland; ca, 100-250 m; Baja California Sur. 

Erythranthe austrolatidens is similar to E. latidens in. its overall aspect but the differences in. 
vestiture and leaf margins and base are readily apparent. These plants have mostly been identified as 
E. floribunda, to which it may be most closely related, but typical E. floribunda occurs all the way 
south in Baja California to the Cape Region. The new species is directly contrasted with both in the 
couplets below. 

1. Vestiture villous-glandular, hairs 0.1-1.0 mm; basal and lower cauline leaves petiolate; margins consistently 
serruiate-mucronulate to serrulate-denticulate, (2-)4-6 teeth per side; fruiting calyces 8-9 mm x 3-5 mm 

Erythranthe austrolatidens 

1. Vestiture stipitate glandular, hairs 0.1-0.3 mm; all leaves sessile; margins entire or barely mueronulate to 
shallowly dentate-mucronuiaie; fruiting calyces 8-12 mm x 4-7 mm Erythranthe latidens 

1 . Medial to distal cauline leaves sessile; stems erect to erect-ascending, mostly branched at the base 

Erythranthe austrolatidens 

1 . All leaves petiolate; erect to decumbent, sometimes procumbent-trailing, branching at proximal to distal 
nodes Erythranthe floribunda 



Nesom: Taxonomy of Erythranthe s 



16. Erythranthe geniculata (Greens) Nesom, Phytoneuron 2011-39: 38. 2011. Mimulus geniculatus 
" Greene. Bull. Calif. Acad, Sci. 1: 280. 1885. Mimulus fioribundusMM. geniculatus (Greene) 
A.L. Grant, Ann. Missouri Bot. Gard. 11: 220. 1924. TYPE: USA. California. Kern Co.: 
Tehachapi, 1884, Mrs. Curran s.n. (holotype: CAS, fragment CAS 290198, fragment MO!; 
isotype: US digital image!). The CAS website provides this information: "Probably the 
holotype was lost in the 1906 fire. This [CAS 290198] is a fragment of that HT, returned by 
Pennell." The MO label (ex CAS) has handwritten "Part of the type." 

Mimulus dudleyi AL. Grant, Ann. Missouri Bot. Gard. 11: 235. 1924. TYPE: USA. California. 
Tulare Co.: rocky cliffs E of the Tule River, 27 Mar 1897, W.R. Dudley s.n. (holotype: DS 
digital image!). 

Annuals, fibrous-rooted or filiform-taprooted. Stems ascending to decumbent or prostrate, 
5-60 cm, simple to diffusely branched. Stems and pedicels moderately villous with multicellular 
eglandular hairs 0.8-2 mm and supitfU-gJjmhilai h.ms 0.1-0.3 mm. Leaves basal and cauline or 
basal mostly deciduous by flowering, blades broadly ovate or elliptic-ovate to triangular, 8-35 x 5-30 
mm, pinnately to subpinnately veined, margins serrate or dentate with 3-10 teeth per side, apex acute 
to obtuse or rounded, base cuneate to rounded or subcordate, petioles 2-10(-35) mm. Fruiting 
calyces campanulate-cylindric, (5-)6-8 mm, weakly inflated, ridged-angled, red-spotted, sparsely to 
moderately villous-glandular, lobes subequal, 1-3 mm, deltate to narrowly triangular or triangular- 
acuminate, usually apiculate or indurate, suberect to spreading-recurving, ciliate. Fruiting pedicels 
12-26(-55) mm. Corollas yellow, red-spotted in throat, spots concentrated or becoming coalescent 
into a somewhat discrete blotch at the base of each of the 3 lower lobes and sometimes the 2 upper as 
well, tube-throats cylindric, 9-11 mm, limbs strongly to weakly bilabiate, expanded 8-14 mm across 
(pressed). Styles glabrous. Anthers glabrous. Capsules obovoid to elliptic, 4-6(-7) mm. 2n = 32 
(reported as Mimulus dudleyi; Heckard 4003, UC). Figs. 5 and 6. Map 8. 

Flowering (Mar-)Apr-Jul. Granite crevices, canyon slopes, and talus, crevices in volcanic 
outcrops (Butte and Tulare cos.), edge of boulders, roadsides, damp sandy soil, sandy water edges, 
gravelly soil and creek bottoms; 200-900(-l 200) m; Calif. 

Only a single collection of Erythranthe geniculata has been seen in the current study from 
each of Butte and Stanislaus counties: Butte Co. : North Table Ml, ca. 7 mi N of Oroville, face of 
basal cliffs on the S side of a small stream, ca. 100 yds S of the fence, ca. 1 mi NW of the parking 
area, cow chute, and Cherokee Road, basalt grassland, 1241 ft, 3 May 2006, Ahart 12,563 (UC). 
Stanisla us Co .: along Hwy 132 ca. 200 yards E of Basso Bridge, 2 mi SW of La Grange, wet soil 
beside pond, 27 May 1969, Allen 355 (DAW). 

Erythranthe geniculata, like E. arenaria, has recently been treated as synonymous with El. 
floribunda. The latter, however, has much smaller corollas and is autogamous, while the flowers of 
E. geniculata and E. arenaria are larger, chasmogamous, and allogamous. in the original description 
of Mimulus geniculatus, Greene noted that it had corollas twice the size of M. floribundus. The 
anther pairs of E. geniculata and E. arenaria are at different levels and the stigma is slightly above 
the upper anther pair, while in E. floribunda both anther pairs and the stigma are at the same level. 

Erythranthe geniculata, E. arenaria, and E. norrisii constitute a group of apparently closely 
related species endemic along the Sierra Nevada. All have ovate-petiolate leaves (only the basal are 
sometimes ovate in E. arenaria) with pinnate to subpinnate venation. The more widespread E. 
floribunda, which is part of the group, also is similar but the three endemics have larger corollas with 
the tube-throats exserted at greater length beyond the calyx margin. 



Nesom: Taxonomy of Erythranthe sect. Mimulosma 25 




Figure 5. Erythranthe geniculata. Granite- Wood)' Rd. Kern Co., California. Photo by Mark Egger, 
3 April 2010. 




Figure 6. Erythranthe geniculata. Indian Wells Canyon. Kern Co., California. Photo by Naomi 
Fraga, 15 April 2011 



Nesom: Taxonomy of Erythranthe s 



17. Erythranthe norrisii (Heckard & Shevock) Nesom. Phytoneuron 2012-39: 39. 2012. Mimulus 

norrisii Heckard & Shevock, Madrono 32: 179. 1985. TYPE: USA. California. Tulare Co.: 
Comb Rocks above Washburn Cove, 2 mi N of Three Rivers, 2800 ft, 1 May 1983, L.L. 
Morris 389 (holotype: .EPS!; isotypes: CAS, FSC, K, MO, NY digital image!, RSA). 

Annuals, fibrous-rooted or filiform-taprooted. Stems ascending to erect-ascending, 2— 15(— 
25) cm, commonly branched from lower nodes. Stems and pedicels villous-glandular. Leaves basal 
and cauline, blades elliptic to elliptic-obovate, 20-35 mm x 10-20 mm, 3-5-palmately veined, 
sometimes with 1-3 distal vein pairs diverging pinnately, surfaces minutely villous-glandular, 
margins subentire to distally denticulate, apex acute to obtuse, base mostly attenuate, petioles 5-10(- 
15) mm. Fruiting calyces campanulate, 4-6 mm, weakly inflated, villous-glandular, sulcate between 
rounded and thickened ribs, lobes subequal, 1.5-2 mm, linear-oblong to oblong-lanceolate with 
rounded to blunt apices, often incurved, villous. Fruiting pedicels 20-35(-50) mm, villous- 
glandular. Corollas yellow with a prominent maroon blotch at the base of each lobe and white patch 
at the 2 sinus bases of the lower lip, weakly bilabiate to regular, sometimes nearly rotate, tube-throats 
cylindric-funnelform, 12-16 mm, limbs bilabiate, expanded 15-30 mm (pressed), lobes oblong- 
obovate to obicular-obovate, apex rounded-truncate. Styles glabrous. Anthers glabrous. Capsules 
narrowly ovoid, 4-6 mm, often slightly exserted In = 32. Map 8. 

Flowering Mar-May. Steep marble outcrops in soil pockets, moss covered marble and 
quartzite ledges, cracks, fractures, and weathered faces, chamise chaparral or blue oak woodland; 
300-1300 m; Calif. (Tulare and Fresno counties). 

Erythranthe norrisii is known only from the Kaweah River drainage and most populations 
are in Sequoia National Park The species is characterized by its short-petiolate leaves with attenuate 
bases, very large corollas with red blotches at the base of each lobe and two white patches on the 
lower lip, very short, purple-dotted calyces with rounded-thickened ribs and with linear-oblong lobes 
incurved in fruit. The capsules often extend beyond the apex of the mature calyces. 

Species of Asia. 

18. Erythranthe stolonifera (Novopokr.) Nesom, Phytoneuron 2012-39: 39. 2012. Mimulus 

stolonifer Novopokr., Bot. Mater. Gerb. Bot. Inst. Komarova Akad. Nauk S.S.S.R. 11: 158 
[with Latin] and 155 [Russian only]. 1949. TYPE. RUSSIA. Province Primoskaja, sinus 
Nachtau [Gulf of Nakhtau], 28 Jun 1911, Dessulavi 1599 (holotype: LE). 

Erythranthe stolonifera (Fig. 6) is endemic to the Ussuri region of Russia (Cape Olympiad, 
Gulf of Nakhtau, Nelka Bay; fide Novopokrovsky 1949), a coastal extension that is essentially 
opposite the islands of Sakhalin (Russia) and Hokkaido (Japan), bordered on the west by northeastern 
China and on the south by North Korea. Plants produce procumbent stems rooting at the nodes and 
arising from a system of lignescent rhizomes. As noted in the protologue, the plants also produce 
distally small-leaved runners from basal cauline nodes. The species is characterized by glandular- 
villous vestiture, essentially ovate, petiolate, pinnately to subpinnately veined leaves with dentate 
margins, long pedicels, and yellow corollas with infundibular tube-throats . It seems likely that its 
closest relative is the North American Erythranthe moschata group (E. moschata, E. moniliformis. E. 
inodora). 



n: Taxonomy of Erythranthe sect. Mimulosma 27 




Figure 7. Collection of Erythranthe stolonifera from Primoski Province, Ri 



Nesom: Taxonomy of Erythranthe s 



Excluded species. 

Erythranthe. bridgesii (Benth.) Nesom (Phytoneuron 2012-n: 0. 2012) was placed by Von 
Bohlen (1995) in the relationship of Mimulus moschatus and M. floribundus, especially based on 
similarities in pollen morphology and perhaps with a tacit assumption that it w ? as related to M. 
moschatus. The placement of E. bridgesii within sect. Mimulosma, however, is problematic, 
especially in view of its glabrous vestiture, strongly palmately veined leaves, and truncate calyx 
margins. The species is tentatively placed as a continentally disjunct member of Erythranthe sect. 
Sinopithecus (Barker et al. 2012), with which it shares glabrous vestiture, sessile, palmately veined 
leaves, calyces with shallowly iobed to subtruncate margins, and broadly spreading, nearly regular 
corolla limbs. 

ACKNOWLEDGEMENTS 

Many thanks to ARIZ, NMC, SD, SRSC, and UC-JEPS for loans of Mimulus sensu lato to 
TEX (where studied), TEX staff for arranging and handling the loans, Amber Schoeneman at TEX for 
the photos of Erythranthe plotocalyx, Phil Tonne at UNM for checking the identity of UNM 
collections filed as Mimulus suksdorfii, Mark Egger and Naomi Fraga for permission to publish 
photos from their web pages, Theo Witsell for company and guidance on a field trip to see 
Erythranthe floribunda in Arkansas, and to DAV, MO, ND-Greene, PH, SMU-BRIT-VDB, TEX-LL, 
UC-JEPS, and UT for help and hospitality during study there. I'm grateful to Naomi Fraga for 
comments on the manuscript and particularly to Matt Carlson sending his dissertation and providing a 
detailed commentary as well as various suggested extensions and modifications. Citations of herbaria 
for duplicates of some types not seen in the present study are from Meinke (1992). This study has 
been supported in part by the Flora of North America Association. 

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Map 1. Erythranthejungermannoides, E. washingtc 
patula. 



E. hymenophylla, E. ampliata, and E. 



Nesom: Taxonomy of Erythranthe sect. Mimulosma 3 1 




Map 2. Erythranthe latidens, E. breviflorus and I hiflaVtla 
Undotted symbols are reports from literature, vouchers not seen 
in present study. 



i: Taxonomy of ErythrantheseA. Mimulosma 22 




o 4. Erythrcmthe moschata and E. moniliformis. Undotted circles are reports from literature., 
vouchers not seen in present study. 




Map 5. Erythranths inodora. 



<onomy of Eryttwaithe sect. / 




Map 6. Erythranthe pulsiferae and E. 
vouchers not seen in present study. 



Undotted symbols are reports from literature, 



ny of Erythran the sect. Mimulosma 35 



Map 7. Distribution of Erythranthe floribunda. Inset show disjunct distribution in Arkansas. Dotted 
circles in Arizona and New Mexico are variant discussed in text. The distribution in Baja California 
Sur continues to the Cape Region. California records from UC-JEPS; Arizona and Mexico records 
are from ARIZ. TEX-LL, and SD. Other records are from various sources, vouchers seen for some 
but not all. 



Nesom: Taxonomy of Erythranthe se 



B Erythranthe norrisii 

(y Erythranthe gen iculata <o^O^ < 



• §:■ 



Map 8. Erythranthe norrisii and /;'. gcniculala.