Phytoneuron

. Leucocarpus (Phrymaceae). Phytoneuron

RECOGNITION AND SYNOPSIS OF MIMULUS
SECT. TROPANTHUS AND SECT. LEUCOCARPUS (PHRYMACEAE)

Guy L. Nesom

2925 Hartwood Drive

Fort Worth, Texas 76109

ABSTRACT

The American genera Hemichaena (5 species) and Leucoc.arp.is (1 species) are formally
brought into Mimulus as Mimulus sect. Tropanthus AL. Grant and Mimulus sect. Leucocarpus (D.
Don) Nesom, comb, et stat. nov. Sect. Leucocarpus includes only M. perfoliatus Kunth. Thieret
(1972) united the five species of Berendtiella and Hemichaena into the single genus Hemichaena and
all five species are formally transferred here to Mimulus. The original Mimulus sect, Tropanthus of
Grant (1924) included only the single species M. treleasei (= Hemichaena levigata) and the section as
revived here and expanded includes these: Mimulus coulteri (A Gray) Nesom, comb, nov.,
Mimulus fruticosus (Benth.) Nesom, comb, nov., Mimulus levigatus (B. Rob. & Greenm.) Nesom,
comb, nov., Mimulus spinulosus (S. Wats.) Nesom, comb, nov., and Mimulus rugosus (Benth.)
Nesom, comb. nov. A lectotype is designated for Hemichaena fruticosa Benth. Sect. Tropanthus and
sect. Leucocarpus are shown by molecular data to have originated as sister to one or the other of the
two main clades of American Mimulus. and for consistency, recognition of Hemichaena and'or
Leucocarpus would require that western American Mimulus be divided into at least two genera. In
contrast to the single-flowered inflorescences characteristic of most Mimulus species, Hemichaena
and Leucocarpus both produce pedunculate multi-flowered bracteolate cymes, which are interpreted
as a primitive feature. The shrubby habit of Hemichaena is paralleled in species of Mimulus sect.
Diplacus and the baccate fruits diagnostic of Leucocarpus are a specialization approached in function
by initially indehiscent capsules in some groups of western US AMtmulus.

KEY WORDS: Hemichaena, Berendtiella, Mimulus sect. Tropanthus, Leucocarpus, Mimulus sect.
Leucocarpus, Phrymaceae

Moiecuiar-phyiogenetic studies of Mimulus (Beardsley and Olmstead 2002; Beardsley et al.
2004; Beardsley & Barker 2005) have provided insights into its phylogenetic structure and putative
relatives. Mimulus sensu stricto, including M. rtngens L., M. alatus Ait, and two Southern
Hemisphere species, is sister to a clade of primarily Australian taxa, including several generally
recognized genera (Fig. 1). Sister to the Australian clade is a branch of two mostly American clades
— Mimulus subg. Synplacus A.L. Grant and Mimulus subg. Schtzoplacus AL. Grant, together
including about 140 species — with the monotypic Phryma L. at their base. Leucocarpus D. Don (1
species) is sister to subg. Synplacus, while Hemichaena Benth. (5 species) is sister to subg.
Schtzoplacus.

In order to retain the name Mimulus for the large number of well-known American species,
an initial stq? is taken here to bring Hemichaena and Leucocarpus into Mimulus. Other actions will
be necessary, since the species identified as Mimulus sensu stricto are sister to the Australian clade.
In case that it proves desirable to maintain the Australian segregate genera, and to maintain Phryma
as a distinct genus, the suggestion by Beardsley and Barker (2005) to conserve the name Mimulus
with a species from within the American lineage is being followed (Nesom and N. Fraga, in prep.).
The much diminished Mimulus sensu stricto will then require a different name. Nesom and Fraga (in
manuscript) also have substantially revised the infragenerie classification of American Mimulus in
conjunction with preparation of the taxonomic treatment of the genus for the Flora of North America
North of Mexico.

. Tropanthus and Leucocarpus 2

Figure 1. Phylogenetic summary
of major groups ofPhrymaceae.

The current concept of Hemichaena was formulated by Thieret (1955, 1972), who
consolidated She species of Berendtiella Wettst. &. Harms and Hemichaena — with the observation
that they are "much alike in habit, habitat, vestiture, foliage, inflorescence, flowers, fruits, and seeds,
all of which convince me that the taxa are congeneric" (1972, p. 89). His observations of their
similarity followed those of Burtt (1965). Argue (1982) found that species of Hemichaena arid
Berendtiella could not be distinguished by pollen morphology, and further support for the close
relationship of Berendtiella and Hemichaena has come from molecular-phylogenetic studies
(Beardsley & Olmstead 2002; Beardsley et al. 2004: Oxelman et al. 2005; Xia et al. 2009).

Pennell (1935) placed Berendtiella along with Mimulus L. and the monotypic Leucocarpus
D. Don in the Scrophulaiiaceae-Gratioleae, "because the flowers and especially the plate-like stigmas
show a clear kinsliip to .Mimulus" (p. 47). In an earlier treatment, however, Pennell (1920) had
constituted the tribe Scrophulariaceae-Mimuieae with Mimulus and Leucocarpus (the treatment
included only Colombian taxa). Burtt (1965) revived the Mimuleae, noting similarities among
Mimulus, Hemichaena, and Leucocarpus in their biiamellate stigmas, campanulate to tubular 5-ribhed
calyces, and loculicidal capsules.

Thieret (1967, p. 93) noted that "Leucocarpus, even though it possesses a baccate fruit, is
obviously closely related to Hemichaena as is evidenced not only by its distinctive stigmas and calyx
but also by its reticulate seeds with intra-reticular lines of a type apparently found nowhere else in the
Scrophulariaceae except in these two genera." Slightly later, Thieret (1972) again recognized that
floral characteristics of Hemichaena and Leucocarpus suggest an alliance with Mimulus, but he
hypothesized on the basis of fruit and seed morphology that Leucocarpus is more distant and placed it
as a relative of the Malaysian Cyrtandromoea Zoll. Interpretations of evidence from floral
morphology are confirmed by the molecular data, wiiich place Leucocarpus as sister to Mimulus
subg. Synplacus — indicating that baccate fruits are a singular specialization evolved within this
group of species that otherwise tiave capsular fruits.

Inflorescences of both Hemichaena and Leucocarpus tend to be bracteolate cymes,
(consistently reduced in two species of Hemichaena to single flowers), while flowers of all species of
American Mimulus are produced singly. The multiple-- flowered inflorescences are distinctive but are
reasonably interpreted as a primitive expression. Taxa closely related to Mimulus-Hemichaena-

r sects. Tropanthus and Leucocarpus 3

Leucocctrpus (Tank et al. 2006; Xia et al. 2009; Albach et al. 2009: Schaferhoff et al. 2010) —
Rehmannia, Triaenophora, Paulownia, Lancea, andMazus — usually produce flowers in racemes or
a thyrse of cymes, sometimes reduced to axillary and solitary flowers in Rehmannia.

Apart from the inflorescence, Hemichaena is not distinct from Mimulus in any consistent
features. Thieret (1972) emphasized the apparent distinction in Hemichaena of superposed
inflorescences, which result from a bud and supernumerary bud in each leaf axil. Supernumerary
buds, however, also occur in M. gemmiparus W.A Weber (the proximal bud becomes a brood bulbil)
andM guttatus Fischer ex DC, (the proximal bud is dormant) (Beardsley 1997; Moody et al. 1999).
The shrubby habit and revolute leaf margins of Hemichaena apparently evolved convergently or in
parallel with species of Mimulus sect. Dipiacus. Beardsley and Olmstead (2002) stated that
"Hemichaena and Berendtiella have united placentae thus making divided placentae a synapomorphy
for Schizoplacus" but Grant (1924) observed that Hemichaena levigata (as Mimulus treleasei, see
next paragraph) is characterized by a separated placenta similar to species of sect. Dipiacus (and all
species of subg. Schizoplacus) — my observations support this for the other species of Hemichaena
as well.

The clearly specialized, baccate fruits of Leucocarpus have a parallel, near-analog in those of
the ten species of Mimulus sect. Oenoe (sensu Thompson 2005), where the capsules are initially
indehiscent, opening along the inner suture after senescence of the plants and after being wet by fall
or winter rains.

Species of Mimulus and Phrymaceae relatives are generally eliaraeterized by five major types
of pollen morphology (Argue 1980, 1983, 1984, 1986; Chadwell et al. 1992). Pollen of both
Hemichaena and Leucocarpus is tricolporate (type II), as is that of Mimulus subg. Synplacus
(excluding sect. Simiolus, which has highly specialized and derivative type I). Pollen of Mimulus
sensu stricto and some of the Australian clade also is tricolporate — Glossostigma has (3)4-6(-7)-
colpate pollen; Barker 1982; A'gue 1986) and Peplidium and Microcarpaea have tricolpate pollen
(Argue 1986). Pollen of Mimulus subg. Schizoplacus (including sect. Dipiacus) is tricolpate or
stephanocolpate (types III, IV, and V). The pollen of Phryma is tricolpate, tectate-perforate to
microreticulate, with simple columellae (type III). Tricolporate pollen is characteristic of close
relatives of Phrymaceae (e.g., Dodartia L., Lancea J.D. Hook. & Thomson, Mazus Lour.; Argue
1984) and it appears to be the primitive type in Phrymaceae.

The morphological similarity of Hemichaena and Leucocarpus to Mimulus has been reflected
in earlier species-level taxonomy. Leucocarpus perfoliatus (Kunth) Benth. was originally described
by Kunth as Mimulus perfoliatus. Hemichaena rugosa (Benth.) Thieret was originally described by
Bentham (1846) as Dipiacus rugosus Benth. Mimulus sect. Tropanthus, as originally described by
Grant (1924), included only the single species M. treleasei AL. Grant, which is a synonym of
Hemichaena levigata (returned to Mimulus as M. levigatus, as treated here). Grant (p. 325) noted that
M. treleasei is "a peculiar species, combining the calyx characters of Eumimulus with the
shrubbiness, the pubescent style, and the separated placentae of Dipiacus." McMinn (1951, p. 114)
observed a resemblance between M. flemingii Munz (= Dipiacus parviflorus Greene) and
Berendtiella levigata "in the configuration of the leaves and flowers" and admitted the possibility that
Dipiacus, which he treated at generic rank, and Berendtiella are derived from a common ancestor.

The geographic distribution of Hemichaena and Leucocarpus is primarily subtropical, but
radiations in traditionally recognized Mimulus have produced species that occur within the
geographic range of Hemichaena — M. pachystylus AL. Grant (endemic to Chiapas; sect.
Paradanthus fide Grant 1924) and M. rupestris Greene (endemic to Morelos; sect. Eryihranthe).

r sects. Tropanthus and Leucocarpus 4

Other species (e.g., M. glabratus Kunth andM guttatus Fischer ex DC.) have ranges that extend from
the western USA through Mexico and Central America to South America.

In summary, Hemichaena and Leucocarpus are both justifiably accomodated as groups
within the bounds of Mimulus — each at the base of one or the other of the two major clades of the
primarily western American species, subg. Schizoplacus and subg. Synplacus. This provides a first
step toward retaining Mimulus as the generic name for these two subgenera, which together' include
about 140 species. If Phryma and Australian segregate genera of Phrymaceae are to be maintained,
the next stq5 is to conserve the name Mimulus with a species from within a western .American group
and adopt an alternative name for the four species of Mimulus sensu stricto.

MIMULUS sect. LEUCOCARPUS (D. Don) Nesom, comb. nov. Leucocarpus D. Don in Sweet, Brit.
Flower Gard. ser. 2, 2: pi. 124. 1831. Type: Conobea alata Graham (= Mimulus pcrfoliatus).

Plants shrubs or suffrutescent perennial herbs, glabrous or subglabrous, eglandular. Stems

0.4-1.8(-2.5) m high, strongly 4-angled to shallowly winged. Leaves thickened, elliptic-oblanceolate
to narrowly lanceolate or narrowly oblanceolate, 9-21(-28) cm x 1.3-4.2(-5.6) cm, margins closely
serrate to serrate-dentate, basally auriculate-clasping and perfoliate. Flowers on short, bracteate
pedicels in axillary, pedunculate cymes of (l-)2-7(-14). Calyces tubular-campanulate, 6-12 mm
long. Corollas 15-22 mm long, deciduous, yellow or white with a yellow throat, bilabiate. Fruits
baccate, 10-18 mm wide, white, with thin skin and with most of the substance of the fruit derived
from the fleshy placenta, septicidally sulcate. indehiscent. Chromosome number apparently not
reported.

A single species.

Mimulus perfoliatus Kunth, Nov. Gen. Sp. (quarto ed.) 2: 371. 1817 [1818]. Leucocarpus
perfoliatus (Kunth) Benth, Prodr. (DC.) 10: 335. 1846. TYPE: Colombia. Crescit in Regno
Novo-Granatensi, no date, Humboldt & Bonpland s.n. (holotype: P).

Conobea alata J. Graham, Edinburgh New Philos. J. 10: 168. 1830. Leucocarpus alatus (J. Graham)
Benth, Brit. Flower Gard. ser. 2, 2: pi. 124. 1833[1831]. TYPE: Mexico. "This plant was
raised in the garden of P. Neill, Esq. at Canonmill [Scotland], from Mexican seeds
communicated by Mr. D. Don as a species of Conobea, and flowered in the greenhouse in
Sept." (from the protologue).

Mimulus perfoliatus ranges from Mexico (Chiapas, Guerrero, Hidalgo, Jalisco. [Michoacan?],
Oaxaca, Puebla, Queretaro, San Luis Potosi, Veracruz) and Central America (Panama, Nicaragua,
Honduras, Guatemala) southward to South America (Bolivia, Colombia, Ecuador, Peru, Venezuela).
It has been collected at elevations of 1500-10,200 feet.

MIMULUS sect. TROPANTHUS AL. Grant, Ann. Mssouri Bot. Gard. 11: 324. 1924. TYPE: Mimulus
treleasei AL. Grant ( :=; Mimulus levigatus).

Hemichaena Benth., PI. Hartw. 78. 1841. TYPE: Hemichaena fruticosa Benth. (= Mimulus
fruticosus).

Berendtia A Gray, Proc. Amer. Acad. Arts 7: 379. 1868 (non Goeppert 1845). Berendtiella Wettst.
& Harms in Engler et Prantl, Nat. Pflanzenfam., Gesamtreg. 2-4: 459. 1899 [a replacement
name for Berendtia A. Gray]. LECTOTYPE (Thieret 1972, p. 92): Berendtia ghiesbrechtii A
Gray (= Mimulus rugosus). Gray did not cite a type for his new genus, in which he included
B. ghiesbrechtii. B. coulteri, and B. rugosa. Thieret cited B. ghiesbrechtii as the type.

s sects. Tropanthus and Leucocarpus 5

Plants perennial, shrubs, stipitate-glandular or glabrous to vernicose. Stems woody, erect.
Leaves pinnately nerved, margins slightly to strongly revolute. Inflorescences axillary cymes,
solitary or superposed, 1-flowered or 2-12-flowered; pedicels shorter to slightly longer than the
calyces. Corollas tubular, yellow to orange or red, sometimes with dotted throat, marcescent,
bilabiate. Anthers glabrous, style pubescent. Capsules dehiscent halfway to completely to the base
along both sutures, placentae separating completely or nearly so. Chromosome number apparently
not reported.

1. Leaves auricu late-clasping; corollas yellow; stamens included Mimuius fruticosus

1. Leaves short-petiolate to subsessile, bases cuneate to attenuate or acute, not clasping; corollas
yellow to orange or red; stamens exserted (slightly so inM levigatus andM. rugosus).

2, Stems and leaves glabrous, vernicose; petiole bases hardly swollen and indurate, blade margins

prominently decurrent as stem wings; corollas orange to yellow Mimuius levigatus

2. At least adaxial leaf surfaces stipitate-glandular; petiole bases swollen and indurate, persistent
after abscission of the blade and upper petiole, blade margins not prominently decurrent; corollas
yellow to orange or red.

3. Leaves 5-22 mm x 1— 3(— 5) mm; young stems densely hirtellous, eglandular, leaves stipitate-
glandular adaxially; inflorescence usually terminal, bracteate; corollas yellow

Mimuius spinulosus

3. Leaves 20-80 mm x 10-35(-50) mm; young stems and both leaf surfaces stipitate-glandular;
inflorescence of axillary flowers arising among large cauline leaves; corollas yellow to orange
or red.

4. Corollas yellow, tube 13-15 mm long, included in or slightly longer than the calyx;

stamens long-exserted Mimuius coulteri

4. Corollas red to orange, tube 25-35 mm long, 2.5-3.5 times longer than the calyx, stamens
slightly exserted Mimuius rugosus

1. Mimuius fruticosus (Benth.) Nesom, comb. nov. Hemichaena fruticosa Benth., PI. Hartw., 78.
1841. Leucocarpus fruticosus (Benth.) Benth., Prodr. (DC.) 10: 336. 1846. LECTOTYPE
(designated here): Guatemala. Prope Quetzaltenango, 1840, K.T. Hartweg 549 (K digital
image!). Other very similar specimens cannot be assumed to be exact duplicates of the type
because of differences in the date or collection number: 1839, Hartweg 549 (LD digital
image!); no date, Hartweg 548 (BM digital image!); 1841, Hartweg 548 (NY digital image!);
1840, Hartweg 548 (PH not seen). The protologue cited only "Quetzaltenango."

Erect herbaceous, suffrutescent, or shrubby plants to 2 m, young stems and leaf surfaces
stipitate-glandular. Leaves sessile, ovate or elliptic, sometimes narrowly so, 4-16 cm long, 1.5-5.5
cm wide, margins coarsely to finely toothed, usually less so or even entire toward the base, base
slightly clasping to cordate-amp lexicaul, sometimes slightly pandurate; petioles 0-1 mm long, bases
swollen and indurate, persistent after abscission of the blade and upper petiole. Inflorescence
axillary, arising among large cauline leaves; peduncles 1-12-flowered, to 4 cm long; pedicels to 20
mm long. Calyces 14-17 mm long. Corollas yellow, 2.5-3.2 cm long, tube 2 times longer than the
calyx. Stamens included, anthers 3 mm long.

Roadsides and road banks, disturbed and open slopes, cliff faces, river banks, sandbars, rocky
places, clay over limestone, oak-pine zones, pine forests, broadleaf cloud forest, evergreen cloud
forest with Quercus, F'inus, Abies, Drimys, Photinia, Clethra, Cornus, and Symplocos; ca. 3000-
11,500 ft: Oaxaca, Chiapas, Guatemala, Honduras, Nicaragua, Costa Rica, Panama.

r sects. Tropanthus and Leucocarpus 6

The name "Hemichaena oaxacana" applied to a collection of Mimulus fruticosus at NY
apparently was never published.

2. Mimulus levigatus (B. Rob. & Greenm.) Nesom, comb. nov. Berendtia levigata B. Rob. &

Greenm., Proc. Amer. Acad. Arts 32: 39. 1897. Hemichaena levigata (B. Rob. & Greenm.)
Thieret, Fieldiana, Bot. 34: 96. 1972. Berendtiella levigata (B. Rob. & Greenm.) Thieret
Ceiba 4: 305. 1955. TYPE: Mexico. Puebla. Calcareous ledges near Tehuacan, 5500 ft, 24
Dec 1895, C.G. Pringle 6294 (holotype: US digital image!; isotypes: A, CAS digital image!,
F digital image!, GH-2 sheets, K digital image!, MO digital image!, PH digital image!, US).
Mimulus treleasei A.L. Grant, Ann. Missouri Bot. Gard. 11: 325. 1924 [1925]. TYPE: Mexico.
Puebla. Tehuacan, 2 Jun 1905, W. Trelease 68 (holotype: MO digital image!).

Erect shrubs to 0.9 m, glabrous, vernicose especially in the younger parts. Leaves opposite
or fascicled, sometimes clustered on spur shoots, elliptic, ovate, or rhombic, sometimes narrowly so,
1.5-5 cm long, 0.4-2.2 cm wide, margins usually revolute, entire to distally serrate with widely
spaced teeth, base acute to attenuate, petioles 0-8 mm long, winged distally, petiole bases neither
strongly swollen nor indurate. Inflorescence axillary, arising among large cauline leaves; peduncles
1-flowered, to 20 mm long; pedicels as peduncles. Calyces 9-15 mm long. Corollas reddish orange,
2.5-3.5 cm long, tube 3 times longer than the calyx. Stamens barely or not exserted, anthers 2-2.5
mm long.

Ledges and rocky soil, limestone slopes, matorral with Morkillia mexicana, Erythroxylon
compactum, Bursera glabrifolia, Sophora secundiflora, Krameria cytisoides, Hintonia latiflora; ca.
5000-6000 ft; Oaxaca, Puebla, Veracruz.

3. Mimulus spinulosus (S. Wats.) Nesom, comb. nov. Berendtia spinulosa S. Wats., Proc. Amer.

Acad. Arts 25: 159. 1890. Hemichaena spinulosa (S. Wats.) Thieret, Fieldiana, Bot. 34: 98.
1972. Berendtiella spinulosa (S. Wats.) Thieret, Cieba 4: 305. 1955. TYPE: Mexico. Nuevo
Leon. Dry limestone cliffs of the Sierra Madre near Monterey, 27 Jun 1888, C.G. Pringle
1952 (holotype: US digital image!; isotypes: F digital image!, GH, K digital image!, USJ).

Erect pendent, or trailing shrubs to 0.9 m, young stems and leaf surfaces stipitate-glandular.
Leaves opposite or fascicled, elliptic, ovate, or obovate; sometimes narrowly so, 5-22 mm long, 1-
3(-5) mm wide, margins entire to several toothed, teeth widely spaced, base acute to attenuate,
petioles 1-2 mm long, bases swollen and indurate, persistent after abscission of the blade and upper
petiole. Inflorescence usually terminal, bracteate; peduncles 1-flowered, to 10 mm long, sometimes
persisting and "thornlike" on dead branches; pedicels as peduncles. Calyces 5-6.5 mm long.
Corollas yellow, tube 13-18 mm long, 1.5-2 times longer than the calyx. Stamens exserted, anthers
0.7-1 mm long.

Outcrops, cliff faces, crevices, talus, slopes, arroyo banks, limestone, gypsum, cemented
gravel, oak-pine forest, scrub-oak zone, oak chaparral with Garrya-Rhus-Agave-Nolina-Pinus or
with Dasylirion-Agave-Cercocarpus-Fraxinus greggii and Pinus, matorral of Acacia, Leucaena,
Pistacia, Vauquelinia, and Dodonea; ca. 2000-5000 ft; Coahuila, Nuevo Leon.

Collections in herbarium TEX-LL that have been recognized by the unpublished name
"Mmulus cebollanus" are within the range of variation of M. spinulosus. Similarly, the name
"Berendtiella pendens" applied to a collection of M. spinulosus at K apparently was never published.

r sects. Tropanthus and Leucocarpus ~j

4. Mimulus coulteri (A. Gray) Nesom, comb. nov. Berendtia coulteri Gray, Proe. Amer. Acad. Arts

7: 380. 1868. Hemichaena coulteri (A Gray) Thieret, Fieldiana, Bot. 34: 94. 1972.

Berendtiella coulteri (Gray) Thieret Ceiba 4: 305. 1955. TYPE: Mexico. No other

information, T. Coulter 1335 (holotype: GH; isotype: K-2 sheets digital images!). A

collection labeled as Coulter 1334 (fide JSTOR) at K apparently is not an isotype.

Erect shrubs to 2 m \oung stems and kaf surfaces stjpriate-gltiulul.u I eaves elliptic, (25-

)30-80 mm x 15— 35(— 50) mm, margins entire to obscurely toothed or undulate, base acute to rounded,

the apex acute to obtuse, petioles 1-2 mm, bases swollen and indurate, persistent after abscission of

the blade and upper petiole. Inflorescence axillary, arising among large cauline leaves; peduncles 1-

5-flowered, to 15 mm long; pedicels to 5 mm long. Calyces 5-8 mm long. Corollas yellow, tube

13-15 mm long, included in or slightly longer than the calyx. Stamens exserted, anthers 1.6-1.8 mm

long.

Rocky or shale to clay or loam slopes, canyons, xerophytic matorral with juniper and oak
deciduous tropical forest, disturbed cloud forest with pine and juniper; ca. 4000-6000 ft.; Guanajuato,
Hidalgo, Queretaro.

5. Mimulus rugosus (Benth.) Nesom, comb. nov. Diplacuz rugosus Benth., Prodr. (DC.) 10: 368.

1846. Hemichaena rugosa (Benth.) Thieret Fieldiana, Bot. 34: 96. 1972. Berendtia rugosa
(Benth.) A Gray, Proc. Amer. Acad. Arts 7: 380. 1868. Berendtiella rugosa (Benth.) Thieret,
Ceiba 4: 305. 1955. TYPE: Mexico. Chiapas. "In Mexico australi in prov. Chiapas," J.J.
Linden 201 (holotype: K, fide Thieret 1972).
Berendtia ghiesbrechtii A. Gray, Proc. Amer. Acad. Arts 7: 380. 1868. TYPE: Mexico. Chiapas.
Ghiesbrecht & Berendt 134 (holotype: GH, fide Thieret 1972).

Erect, arching, or sometimes pendent shrubs to 4 m, young stems and both leaf surfaces
stipitate-glandular. Leaves opposite or fascicled, ovate or elliptic, sometimes narrowly so, 20-70 mm
long, 10-35 mm wide, margins coarsely to finely many to few toothed, the base cimeate to attenuate,
petioles 1-7 mm long, bases swollen and indurate, persistent after abscission of the blade and upper
petiole. Inflorescence axillary, arising among large cauline leaves; peduncles 1-3-flowered, to 15
mm long; pedicels to 12 mm long. Calyces 8-14 mm long. Corollas orange to red, tube 25-35 mm
long, 2.5-3.5 times longer than the calyx. Stamens exserted, anthers 1.5-2 mm long.

Rocky slopes, ridges, cliff faces, roadsides and roadbanks, matorral, oak and pine-oak-
juniper-arbutus woodlands; ca. 4600-10,000 it; Chiapas, Guatemala, Honduras.

ACKNOWLEDGEMENTS

I'm grateful to the staff of TEX-LL for help in obtaining literature and loans of specimens and
to Barney Lipscomb at BRIT for help with literature. This study was done in conjunction with
preparation of the FNANM treatment of Mimulus and supported by the Flora of North America
Association.

LITERATURE CITED

Albach, D.C, K. Yan,, S.R Jensen, and H.Q. Li. 2009. Phylogenetic placement of Triaenophora

(formerly Scrophulariaceae) with some implications for the phylogeny of Lamiales. Taxon

58:749-756.
Argue, C.L. 1980. Pollen morphology in the genus Mimulus (Scrophulariaceae) and its taxonomic

significance. Amer. J. Bot. 67: 68-87.
Argue, C.L. 1983. A biometric and taxonomic study of pollen character variation in Berendtiella and

Hemichaena (Scrophulariaceae). Canad. J. Bot. 61: 53-62.

r sects. Tropanthus and Leucocarpus 8

Argus, C.L. 1984. Pollen morphology in Dodartia, Lancea, Leucocarpus, and Mazus and an
analysis of pollen morphotypes in the Mimuleae (Scrophulariaceae). Canad. J. Bot. 62:
1287-1297.

Argue, C.L. 1986. Pollen morphology of Amphianthus, Artanema, Curanga, Glossostigma, and
Peplidium (Scrophulariaceae - Gratioleae). Amer. J. Bot. 73: 1570-1576.

Barker, W.R. 1982. Evolution and biogeography of arid Australian Scrophulariaceae. Pp. 341-350,

in W.R. Barker andP.J.M. Greenslade (eds.), Evolution of the Flora and Fauna of Arid

Australia. Peacock Publications, Adelaide, South Australia.
Beardsley, P.M. 1997. Colorado's rare endemic plant, Mimulus gemmiparus, and its unique mode of

rq^roduction. M.S. thesis (Botany), Colorado State Univ., Ft. Collins.
Beardsley, P.M. and R.G. Olmstead. 2002. Redefining Phrymaceae: The placement of Mimulus,

tribe Mmuleae, and Phryma. Amer. J. Bot. 89: 1093-1 102.
Beardsley, P.M., S.E. Schoenig, IB. Whittall, and R.G. Olmstead. 2004. Patterns of evolution in

western North American Mimulus (Phrymaceae). Amer. J. Bot. 91: 474-489.
Beardsley, P.M. and W.R. Barker. 2005. Patterns of evolution in Australian Mimulus and related

genera (Phrymaceae similar to Scrophulariaceae): A molecular phylogeny using chloroplast

and nuclear sequence data. Austral. Syst. Bot. 18: 61-73.
Bentham, G. 1846. Scrophulariaceae. Prodr. (DC.) 10: 186-586. Diplacus (p. 368), Mimulus (pp.

368-373), Eunanus (p. 374).
Burtt, B.L. 1965. The transfer of Cyrtandromoea from Gesneriaceae to Scrophulariaceae, with notes

on the classification of that family. Bull. Bot. Survey India 7: 73-88.
Chadwell, T.B., S.J Wagstaff, and P.D Cantino. 1992. Pollen morphology of Phryma and some

putative relatives. Syst. Bot. 17:210-219.
Grant, A.L. 1924. A monograph of the genus Mimulus. Ann. Mssouri Bot. Gard. 11: 99-388.
McMinn, H.E. 1951. Studies in the genus Diplacus (Scrophulariaceae). Madrono 11:33-128.
Moody, A, P.K. Diggle, and D.A Steingraeber. 1999. Developmental analysis of the evolutionary

origin of vegetative propagules in Mimulus gemmiparus. Amer. J. Bot. 86: 1512-1522.
Oxelman, B., P. Kornhall, RG. Olmstead, and B. Bremer. 2005. Further disintegration of

Scrophulariaceae. Taxon 54: 411^125.
Pennell, F.W. 1920. Scrophulariaceae of Colombia. Proc. Acad. Nat. Sci. Philadelphia 72: 136-188.
Pennell, F.W. 1935. The Scrophulariaceae of eastern temperate North America. Proc. Acad. Nat.

Sci. Philadelphia 1: 1-650. Mimulus Linnaeus (pp. 112-136).
Schaferhoff, B., A Fleischmann, E. Fischer, D.C. Albach, T. Borsch, G. Heubl, and K.F. Muller.

2010. Towards resolving Lamiales relationships: Insights from rapidly evolving chloroplast

sequences. BMC Evol. Biol. 10: 352 (22 pages).
Tank, D.C, P.M Beardsley, S.A Kelchner, and R.G. Olmstead. 2006. L.AS. Johnson Review No.

7. Review of the systematics of Scrophulariaceae s.l. and its current disposition. Austral.

Syst. Bot. 19: 289-307.
Thieret, J.W. 1954. The tribes and genera of Central American Scrophulariaceae. Ceiba 4: 164-184.
Thieret, J.W. 1955. The status of Berendtia A Gray. Ceiba 4: 304-305.

Thieret, J.W. 1967. Supraspecific classification in the Scrophulariaceae: A review. Sida 3: 87-106.
Thieret, J.W. 1972. Synopsis of Hemichaena, including Berendiiella (Scrophulariaceae).

Xia, Z., Y.-Z. Wang, and J.F. Smith. 2009. Familial placement and relations of Rehmannia and

Triaenophora (Scrophulariaceae s.l.) inferred from five gene regions, Amer. J. Bot. 96: 519-