KATHLEEN BURT-UTLEY and JOHN E UTLEY

Institute for Systematic Botany

Department of Cell Biology, Microbiology, and Molecular Biology

University of South Florida

Tampa, Florida 33620-5150

kburtutl@uno.edu; jutley@uno.edu

ABSTRACT

Ten new species of Begonia are described, discussed, and illustrated: Begonia wilburi Burt-
Utley & Utley, Begonia gentryi Burt-Utley & Utley, Begonia liesneri Burt-Utley & Utley, Begonia
mcphersonii Burt-Utley & Utley, Begonia pseudopeltata Burt-Utley & Utley, Begonia
aguabuenensis Burt-Utley & Utley, Begonia sukutensis Burt-Utley & Utley, Begonia panamensis
Burt-Utley & Utley, Begonia gracilioides Burt-Utley & Utley, and Begonia tenuis Burt-Utley &
Utley. Begonia militaris L.B. Sm. & E.G. Schub. is evaluated and B. sciadophora L.B. Sm. & B.G.
Schub. is synonymized with it while B. pustulata Liebm. and B. ludicra A DC. are recognized as
species endemic to Mexico.

KEY WORDS: Begoniaceae, Begonia. Mexico, Central .America, Colombia

Continuing research with the Begoniaceae and preparation of the Begoniaceae for the Flora
Mesoamericana has resulted in the recognition of a number of new species (Burt-Utley & Utley 1999;
2011), including the 10 that are described herein. One of these species, B. wilburi, is unique in
Central America because of its potential dioecy and is only one of two species in the Flora with
turbinate capsules. The other known obligate dioecious species in Mexico and Central America are
from Mexico, B. extranea L.B. Sm. & B.G. Schub. (Guerrero, Mexico, Michoacan, and Jalisco), and
B. nemoralis L.B. Sm. & B.G. Schub. (Michoacan), both of which are tuberous. .'Another tuberous
species from Mexico and Central America, B. biserrata Lindl., is either monoecious or dioecious
(Burt-Utley & McVaugh 2001). Two other species also stand apart from other Begonia in Chiapas
and Central America, B. gracilioides and B. tenuis, because they are tuberous and also produce bulbils
in their leaf axils that can then develop into new individuals. In Central America, similar bulbils have
been observed only on B. weberlingii Irmsch. and B. ignea (Kl.) A DC. In contrast, in Mexico, 17 of
the 37 known tuberous taxa are known to produce bulbils and are most common in central and
western Mexico (Burt-Utley, pers. obs.). The species described represent diverse phenetic groups
including sect. Casparya (Kl.) A DC, sect. Gireoudia (Kl.) A DC, sect. Weilbachia (Kl. & Oerst.)
A DC, and sect. Knesebeckia (Kl.) A. DC.

1. BEGONIA WILBURI Burt-Utley and Utley, sp. nov. TYPE. COSTARICA. San Jose. Ca 26 km

S of La Georgina on Interamerican Hwy to San Isidro de El General, 1600-1800 m, 19 Oct

1974, J. Utley & K. Utley MW $ (holotype: DUKE; isotypes: CR, USF). Figure 1.

Suffrutescent herbs to at least 1.5 m, frequently branching, potentially dioecious, upper
internodes slender, 3.5-7.5 cm, 1.5-4.5(-6) mm diam, densely hirtellous to tomentose with
short ferrugineous villi to 0.4 mm. Stipules persistent to deciduous, asymmetric, unequal, the larger
appearing hemi-ovate to hemi- orbicular with an oblique acute apex, 4-5.5 x 8-11 mm, the smaller,
oblong to triangular, 1.2-8 x 1-4 mm, apically acute, marginally ciliate and denticulate to serrulate,
hirtellous above and beneath with villi to 0.3 mm; petioles 0.7-5.4 cm, densely hirtellous with villi to
0.5 mm; leaf blades oblique or rarely straight, asymmetrically elliptic to oblong, (5.5-)8.5-18+ x

(3.6-)4.8-8.1 cm, basally unequal-sided, apically attenuate-acuminate, marginally doubly ciliate-
crenate to doubly ciliate-serrate and serrulate, above and below densely hirtellous; (6-)ll to 14
pinnately nerved on the broad side of the blade. Staminate inflorescences appearing
subumbelliferous with perhaps occasionally than 2 branches at the lowermost node, 3-10 or more-
flowered; peduncles 1-6.7 cm, densely hirtellous with villi to 1 mm; bracts deciduous, elliptic, to 6.5-
11 x 3.5-5.5 mm, marginally ciliate-serrulate to ciliate-laciniate. Pistillate inflorescences 1-
flowered; peduncles (1.5— )3. 5-4.5 cm. densely hirtellous with villi to 1 mm: bracts minute, 1^ mm,
apically lobed. Staminate flowers with pedicels 6-13 mm, hirtellous; sepals 2, elliptic to ovate, 8.5-
13 x 8-10 mm, marginally occasionally ciliate distally, glabrous to pilose, translucent white to white
sufficed pink; petals 2, narrowly obovate, 7.5-8.5 x 3.5-5.5 mm, translucent white or white mottled
pink; stamens 40-59, appearing monadelphous; filaments 1-1.5 mm; anthers narrowly oblong to
obovate, 1.5-2 x 0.4-0.6(-0.8) mm. Pistillate flowers with pedicels l-A mm, densely hirtellous;
bracteoles wanting; tq^als 5, subequal, ovate, 12.5-15 x 4-10 mm, pilose without, white or white
suffused pink; ovaries trilocular with bipartite placentae, 11 mm, densely hirtellous; styles multi-
branched to 2 mm; stigmas at tips of stylar branches. Capsules with pedicels (5— )8— 14 mm; bodies
turbinate, unbeaked, 15-20 mm; wings or horns 3, subequal, oblong to obovate, marginally entire to
undulate or denticulate distally.

Etymology. It is a great pleasure to name this distinctive species in honor of Dr. Robert L.
Wilbur at Duke University who collected extensively in Costa Rica and Panama and has been
particularly dedicated to their montane floras. Dr. Wilbur devoted much of his career to developing
the fine vascular plant collection in the Duke Herbarium. He was a fine dissertation advisor and
mentor for my research with Begoniaceae.

Distribution and habitat. Begonia wilburi occurs at elevations between 1600 m and 3350 m
in montane cloud forests and wet secondary forests on steep mountainsides in the Cordillera
Talamancas with Alnus and other vegetation in Costa Rica and western Panama.

Begonia wilburi is characterized by its apparent dioecy and 1-flowered pistillate
inflorescences, immediately standing apart from other known Mesoamerican taxa, except
occasionally B. biserrata in its dioecy (Burt-Utley &. McVaugh 2001). While it is unclear if B.
wilburi is dioecious, branches of specimens examined either had only staminate inflorescences or
pistillate inflorescences at each node, and no branches had both staminate and pistillate
inflorescences. In all species studied from Mexico and Central America, only the tuberous B.
cebadillensis Houghton ex L.B. Sm. & B.G. Schub. was observed with separate staminate and
pistillate inflorescences on the same stem or on the same individual (Burt-Utley pers. obs.). Although
there are a. number of collections of B. wilburi. only two examined {Picado & Gamboa 251, MO;
Utley & Burt-Utley 9033, USF) consist of a branch with staminate inflorescence and a branch with
pistillate material, and it is unclear if they came from the same individual. It is not surprising that
only two collections of B. wilburi have both staminate and pistillate material because staminate stems
or individuals of this species are inconspicuous when flowering, and this is even more true of
pistillate stems with their short peduncles with one flower (Burt-Utley pers. obs.).

Begonia wilburi was first collected by Adolph Tonduz in April 1898 ("Herb. Instit. physico-
geogr. nat. costaricensis" #12274) and subsequently annotated by C. de Candolle as B. cooperi C.
DC, a vegetatively similar species. Begonia wilburi, however, appears dioecious with "horned" or
turbinate capsules and ovaries, an unusual character that immediately places it in the predominantly
South American section Casparya (Kl.) Warb. Within Central America, the remaining Begonia
species have ovaries, capsules, or rarely berries with wings and are all monoecious (pers. obs.). The
only other species in this section known from Costa Rica and Panama is B. urticae L. f. and B.
wilburi is easily separated from, it by vegetative and floral characters, including its larger leaf blades

Burt-Utiey and Utiey: New species and n

lsotype (Utiey & Utiey 1470, USF).

[C5.5-58.5-18 x (3.6-)4.8-8 ci
8-10 mm vs. 3-5 x 3-4 mm),

i notes on Begonia 4

Begonia wilburi appears most closely related to the Colombian endemic, B. ursina L.B. Sm.
& B.G. Schub., described from fragments, which also is presumably dioecious, with similar few-
flowered male inflorescences and one-flowered female inflorescences on separate stems. Moreover,
both species are reported from a high elevations (3100 m) and have hirsute to hirtellous stems, leaves,
sepals, petals, tepals, and ovaries (pers. obs.; L.B. Sm. & B.G. Schub. 1946). Leaf blades of B.
wilburi are larger than those of B. ursina [(5.5-)8.5-18+ x (3.6-)4.8-8. 1 cm vs. 5-5.7 x 1.6-2.4 cm].
Stipules also differ in form between these two species, with those of B. wilburi having two different
shapes at each node (the larger hemiorbicular and the smaller oblong to triangular) while those of B.
ursina are ovate. Bracts are large and elliptic in staminate inflorescences (6.5-11 x 3.5-5.5 mm) of B.
wilburi but much smaller to minute (1^1 mm) in their pistillate inflorescences. In B. ursina,
staminate bracts are described as ovate (10 mm), and apparently are equal in size to the "bracteoles"
of its pistillate flowers. It seems very likely that the authors actually were describing the pistillate
bracts of B. ursina rather than bracteoles. Begonia wilburi is also distinguished from B. ursina in its
subequal 5-tepaled pistillate flowers, in contrast to the 6-tepaled pistillate flowers of B. ursina with
three subequal outer tepals and three smaller inner subequal tepals. Six-tepaled pistillate flowers are
not unique to B. ursina and have been observed in other species in sect. Casparya (L.B. Sm. & B.G.
Schub. 1946). While both species have unbeaked capsules, the horns of B. wilburi are straight, broad,
and oblong to obovate, in contrast to those of B. ursina which are narrow and falcate.

Specimens examined. COSTA RICA. San Jose. Bords du Rio de la Mala, Via au Copey
Apr 1898, Tonduz 12274 3 (CR); Canton Perez Zeledon, P.N. Chirripo, Cordillera de Talamanca,
sendero al Mirador, 9°33'20' N, 83°40'15" W, 18 Aug 1995, Picado & Gamboa 251 (3 & 2 MO, 3
USF); Canton Perez Zeledon, 1 km de Division a la par de Carr. Interamericana, 9 Dec 1996, Hammel
& Model 20575 (MO); slopes of Cordillera de Talamanca near la Division, N of San Isidro de El
General, 6 Feb 1963, Williams, Jimenez M., & Williams 24385 3 (F). 10.8 km S of La Georgina on
Interamerican Hwy to San Isidro, 25 Jun 1995, 8000 ft, Utley & Utley 9031 3 (USF); 9.8 km S of La
Georgina, 25 Jun 1995, 8000 ft, Utley & Utley 9033 (3 MO, US. 3 & ? USF). Puntarenas. N of
San Isidro del General, 12 Aug 1971, 7000-11000 ft, Vaughan, Dwyer; Spellman, & Wunderlin 679 3
(MO). PANAMA. Chiriqui. Vic. of Cerro Punta, 0.5 mi SE of Entre Rios, 1 mi by rd from town of
Cerro Punta, 25 Nov 1979, 2000 m, Croat 48573 $ (MO); Volcan Barii (E slope), deep draw W of
Finca Yen, 17 Mar 1979, 8000 ft, Hammel, D 'Any, &Averett 6452 (MO).

2. BEGONIA GENTRYI Burt-Utley & Utley, sp. nov. TYPE. COLOMBIA Choco. Alto de Buey
500-1200 m, 8 Jan 1973, A. Gentry & E. Forero 7262 (holotype MO; isotypes MO, US).
Figure 2.

Rhizomatous herbs; internodes very short, to 9 mm, 7-9.5 mm diam, glandular and villous-
squamose with trichomes 2-4(-6) mm. Stipules apparently persistent, narrowly triangular, 13-21 x
3-9 mm, marginally entire, villous, strongly keeled, with the keel fimbriate- laciniate; petioles 9-21
cm, villous with stout trichomes 2-5 mm intermixed with minute glandular hairs; leaf blades
asymmetric, oblique to transversely elliptic, reniform or ovate, 12-28 x 8.5-19 cm, basally cordate,
apicaily acute to short acuminate, marginal!} ecihate and somewhat undulate, glabrous to minutely
glandular above and glandular to very sparingly villous below, especially on nerves, with trichomes
0.3-2 mm; 10-12-palmatinerved. Inflorescences asymmetrically cymose, greatly exceeding the
foliage, ca 14-flowered; peduncles 40-70 cm, sparingly villous with trichomes 1.5-6 mm; bracts
caducous, oblong-elliptic, 20 x 11-12 mm, apicaily truncate, marginally entire. Staminate flowers
with pedicels ( 1 1— ) 1 5—20 mm, glabrous; sepals 2, suborbicular to broadly oblong, 11-14 x 13-14
mm, glabrous, light orange; petals 2, obovate, 10-12 x 7 mm; stamens 25-32; filaments appearing
free, less than 0.1 mm; anthers elliptic to obovate, 2-2.2 x 0.4-0.6 mm. Pistillate flowers with
pedicels 12-13 mm, glabrous; bracteoles wanting: sepals 2, transversely elliptic, 11-15 x 10-14 mm,
glabrous, light orange; petals 2, obovate, 10 x 8 mm; ovary trilocular with bipartite placentae, 6-6.5

Burt-Utley andUtley: t

Figure 2. Begonia, gentryi. Hololype (Gentry & Forero 7262, MO).

mm, glabrous to sparingly glandular; styles 1.7-2 mm, free to the base: stigmas bicornute. Capsules
with pedicels 19-27 mm; bodies 9-11 mm; locule chambers externally appearing broadly ovate, 7.5-
8.5 x 6.5-8 mm; wings 3, unequal, the largest wing ovate to elliptic, 19-23 x (9-)13-18 mm, apically
rounded to subacute, the other 2 marginiform and triangular to lunate.

Etymology. Begonia gentryi is named in honor of the late Dr. Alwyn H. Gentry (1945-
1993), who was a Curator at the Missouri Botanical Garden before his untimely death in an airplane
crash in Ecuador.

Distribution and habitat. Begonia gentryi is known from the Choco in Colombia in tropical
wet forests at elevations between 500 and 1200 m and has been collected in the Darien in Panama.
According to Whiteford and Eddy, B. gentryi is common above 900 m in the Serrania de Jungorodo in
Panama.

Begonia gentryi is distinguished by its rhizomatous habit, eciliate leaf margins, and pistillate
flowers with two sepals and two petals. It is one of about four rhizomatous species occurring in
Colombia and is readily distinguished from them by its staminate and pistillate flowers consistently
with two sq^als and two petals, in contrast to two of the remaining three species that have apetalous
flowers (B. nelumbiifolia Schlecht. & Cham, and B. urophylla Hook.) and B. sericoneura Liebm. with
apetalous staminate flowers but pistillate flowers that are either apetalous or have a single petal (Burt-
Utley 1985). Begonia gentryi stands apart from known Mesoarnerican rhizomatous taxa in its orange
pistillate and staminate Howlers and, with the exception of B. mucronistipula C. DC, pistillate flowers
with two sepals and two petals. Flowers of species from the region of the Flora characteristically
have sepals, petals, or tqials that range from white to deep pink or rarely yellow-green (Burt-Utley
pers. obs.). Begonia gentryi is also notable for the small but conspicuous cystospheres, which arG
evident on all plant parts except the petals and roots. It appears most closely related to the
Panamanian endemic, B. mucronistipula, with which it shares its rhizomatous habit and similar
pistillate flowers (Burt-Utley & Utley 2011). Other rhizomatous Central American taxa have
pistillate flowers that are apetalous or have one petal or five subequal tepals. Begonia gentryi is
readily distinguished from B. mucronistipula by a number of characters, including its internodes with
a villous-squamose indument (vs. glabrous), villous stipules and petioles (vs. glabrous), larger anthers
(2-2.5 mm vs. 1-1. 6 mm), and its" large primary wings [19-23 x (9-)13- 18 mm vs. (11-)15-17 x 7-
9.5 mm]. Begonia gentryi also occurs in tropical wet forests at a lower elevations than those reported
for the montane B. mucronostipula (500-1200 m vs. 1500-2100 m). Although B. gentryi shares
character's with B. mucronostipula, which was tentatively included in sect Gireoudia (Doorenbos et
al. 1998), the sectional affiliations of both species are unclear (pers. obs.).

Additional specimens examined. PANAMA. Darien. Serrania de jungorodo, Mamey, 900
m, 10 Mar 1982, Whiteford & Eddy 460 (BM). COLOMBIA Choco. S ridge of Cerro Mecana,
6°16'N, 77°18'W, 500-700 m 7 Jan 1984, Juncosa 1758 (MO, USF); Rio Mutata ca 3 km above its
junction with Rio El Valle NW of Alto de Buey, 850 m, 7 Feb 1971, Lellinger & de la Sota 182 (US).

3. BEGONIA LIESNERI Burt-Utley & Utley, sp. nov. TYPE. COSTA RICA. Cartago. Cerro
Doan, 3 km E of Cachi, 23 Apr 1969, R.W. Lent 1601 (holotype: F; isotypes: NY, US).
Figure 3.

Suffrutescent herbs to 80 cm tall; upper internodes (1.5-)4-6 cm, 2-4 mm diam, glabrous,
cystospheres abundant. Stipules deciduous, asymmetrically ovate to oblong-ovate, 15-18 x 7-8 mm,
marginally entire, glabrous, keeled, with only the keel villous; petioles 1.5-3.5 cm, glabrous to very
sparingly villous, with the stout villi 0.1-0.7 mm; leaf blades straight to weakly arcuate,
ally narrowly elliptic to ovate, 7-13 x 1.8-4.3 cm, basally unequally sided, apically

attenuate-acuminate, marginally ciliate-serrulate and ciliate-serrate, glabrous throughout above,
villous beneath only on principal nerves with stout villi to 0.7 mm; (10— )12— 15 pinnately nerved on
the broad side of the blade. Inflorescences shorter than the leaves, laxly cymose, ca +/- 7-flowered;
peduncles 3.5-6.5 cm, glabrous to villous with few villi to 0.7 mm; bracts deciduous, ovate to
obovate, 9-12 x 5-7 mm, marginally serrulate, glabrous to sparingly villous, keeled, or the keel only
villous. Staminate flowers with pedicels 10-15 mm, sparingly short villous; sepals 2, ovate to
oblong or elliptic, 9-16 x 6-9.5 mm, glabrous to glandular and sparingly short villous; petals 2,
obovate, 9-13.5 x 4.5-6.5 mm, distally shallowly lobed, glabrous; stamens +/- 23, appearing
somewhat monadelphous; filaments 0.8-1.3 mm; anthers narrowly obovate to elliptic, 1.8-2 x 0.5-
0.7 mm. Pistillate flowers with pedicels to 15 mm, villous; bracteoles present, deciduous, obovate,
8-9 x 6-7 mm, apically cilia te-laciniate, marginally ciliate-serrulate, sparingly villous; sq>als at
anthesis unknown; petals/tepals unknown: ovary trilocular with bipartite placentae, 8-10 mm,
hirtellous and minutely glandular; styles and stigmas unknown. Capsules with pedicels to 24 mm;
bodies 11-14 mm; externally locule chambers elliptic, 8-11 x 6-7 nun; wings subequal to unequal,
the largest one asymmetrically triangular, 5-12 x 11-15 mm, the others asymmetrically triangular.

Etymology. Begonia liesneri is named in honor of Mr. Ronald Liesner at the Missouri
Botanical Garden, who first recognized this as a new species.

Distribution and habitat. Begonia liesneri is known only from Cartago Province where it
occurs in cloud forests or rainforests at bases of cliffs between 1400 and 1450 m.

Begonia liesneri is characterized by a suite of characters including its suffrutescent habit,
glabrous internodes, pinnately nerved leaf blades, large bracteoles, ca 7-flowered inflorescences, and
large capsules with subequal to unequal capsule wings. With its several-flowered inflorescences that
are shorter man the foliage, B. liesneri is potentially an inconspicuous component of the vegetation
and easily overlooked by collectors. Without pistillate flowers available, it is difficult to determine
the relationships of B. liesneri with other Central American taxa because caulescent species with
similar staminate flowers could have pistillate flowers with two sepals and one or two petals or four
to five subequal tepals (Burt-Utley, pers. obs.). Begonia liesneri is distinguished from suffrutescent,
pinnately nerved Begonia species found in Costa Rica by its several-flowered inflorescences, large,
marginally serrulate bracts, and, in all but B. tonduzii C. DC, its subequal capsule wings. With the
exceptions of B, tonduzii, B. wilburi (described herein), and the Costa Rican endemic B. cooperi C.
DC, it is the only other pinnately nerved, pubescent, suffrutescent taxon from Central America (pers.
obs.). From B. tonduzii it is readily distinguished by its larger bracts [9-12 x 5-7 mm vs. 1 . 3—3 (—5 ) x
1-3(^1) mm], larger staminate sepals (9-16 x 5-10.5 mm vs. 5-10.5 x 4.5-9 mm), and larger
capsules (11-14 mm vs. 6-11 mm) (Burt-Utley, pers. obs.). It differs from B. cooperi C DC in its
much larger staminate sepals [9-16 x 6-9.5 mm vs. (3-)4-6(-8) x 3.5-5(-8) mm], large,
conspicuously bracteolate pistillate flowers (vs. ebracteolate). persistent petals, and larger capsules
[11-14 mm vs. (4-)6-8.5 (-9.5) mm] (Burt-Utley, pers. obs.). The only other pinnately nerved
suffrutescent species bearing several flowers is B. wilburi, but B. liesneri differs immediately from
this potentially dioecious species in its monoecy and non-turbinate capsules.

Additional specimens examined. COSTARICA. Cartago. Tapanti, Orosi, 1400 m, 29 Jan
1983, /. A. Chacon 259 (USF).

Surt-Utiey and Utiey: New species and notes on Begonia

Figure 3. Begonia liesneri. Holotype (Lent 1601, F).

4. BEGONIA MCPHERSONII Burt-Utley & Utley, sp. nov. TYPE. PANAMA. Bocas del Toro.
Vic. of Cerro Colorado mine above SanFelix, trails N of road on continental divide. 8°35'N,
81°50'W, 1500 m, 26 Jan 1988, G McPherson 12019 (holotype: MO; isotypes: PMA, USF).
Figure 4.

Rhizomatous herbs; internodes 0.5-1.6 cm, 0.4-1.3 cm diam, lanate with fine sericeous
trichomes 1.5-5 mm. Stipules persistent, coriaceous, asymmetrically ovate, 1.2-2.7 x 1-1.1+ cm,
marginally entire, glabrous keeled, or the keel only pilose, cystospheres abundant; petioles (10.5— )13—
23 cm, tomentose with fine sericeous villi 1.5-4 mm; leaf blades oblique to transverse, symmetric to
asymmetric, elliptic to oblong in outline, 10.5-18 x 8-17.5 cm, basally cordate, apically with no
distinct apex, marginally ciliate-serrate and ciliate-serrulate, deeply asymmetrically palmately lobed
usually with 4 to 5 major attenuate-acuminate lobes to Vi the blade length, sparingly pilose above
especially above the petiole-blade junction, lanate on primary nerves below but pubescence less dense
in intercostal regions; 10-11-palmatinerved. Inflorescences greatly exceeding the foliage, weakly
asymmetric, densely to laxly cymose with elongate branches, many-flowered; peduncles (21-)30-49
cm, lanate to pilose; bracts caducous, the lowermost apparently completely encircling the inner in
bud. subequal, broadly navicular, broadly ovate, 1.7 x 1.4 cm, pilose, cystospheres abundant.
Stamioate flowers with pedicels 9-10.5 mm, pilose; sepals ovate, 5-7 x 7-9 mm, pilose, pink-white;
petals 0-2, narrowly obovate to ovate, 4.5-6 x 1.2-1.5 mm; stamens 23-35; filaments 0.5-0.8 mm,
on a raised torus and appearing somewhat monadeiphous; anthers obovate to elliptic, 1—1.4 x 0.4-0.5
mm. Pistillate flowers with pedicels 6-8 mm, pilose; bracteoles wanting; sepals suborbicular to
transversely broadly elliptic, (5.5-)8-9 x 8-9.5 mm, pilose, pink-white; petals 0-1, obovate, 6 x 2,5-3
mm; ovary trilocular with bipartite placentae, 3.5-4.5 mm, pilose, cystospheres present; styles 3, 1.5-
2 mm, connate over Vi their length; stigmas bicornnte. Capsules with pedicels 13-19 mm; bodies 6-
7.5 mm with conspicuous cystospheres; locule chambers externally ovate, 5-6.5 x 3.5-5.5 mm; wings
3, unequal, the largest wing asymmetrically elliptic, 11-14.5 x 8-10.5 mm, the second one
asymmetrically triangular, 5-7 x 5-6 mm and the third marginiform.

Etymology. It is a pleasure to name this species in honor of Dr. Gordon McPherson at the
Mssouri Botanical Garden, who collected this species. Gordon was our field companion and guide in
Panama when we collected Begonia and Bromeliaceae in 1988.

Distribution. Begonia mcphersonii is known only from the provinces of Bocas del Toro and
Chiriqui in Panama near the continental divide between (800-)1200-1500 m.

Begonia mcphersonii appears most similar and closely related to other two species in sect.
Gireoudia (Kl.) A. DC, B. involucrata Liebm. and B. broussonetiifolia A. DC, because of its deeply
lobed leaves, tomentose indument, and its lowermost bracts with the outer apparently encircling the
inner. This latter character is unusual among Central .American Begonia and has only been observed
in the suffrutescent taxa in sect. Gireoudia, B. involucrata and B. broussonetiifolia A DC. (Burt-
Utley 1985). From B. broussonetiifolia, B. mcphersonii is readily distinguished by its rhizomatous
habit, shorter and stouter internodes [0.5-1.6 cm, 0.4-1.3 cm diam vs. (0.9)1. 8-4.4 cm, (2-)3-6 mm
diam], palmately multilobed blades (4-5 major lobes vs. 1-2 lobes), and longer peduncles [(21-)30-
49 cm vs. (14.6-)17.5-27.5(-29) cm. Begonia mcphersonii stands apart from B. involucrata in a
number of characters including its habit and elongate inflorescence internodes. Upper branches and
frequently the lower branches of B. involucrata inflorescences are greatly reduced in length, resulting
in inflorescences that are strongly congested distal ly (Burt-Utley 1985). Because of these differences
in branch length, there are conspicuous differences in pedicel length between these two species.
Pedicels of staminate flowers, pistillate flowers, and capsules of B. mcphersonii are much shorter than
those of B. involucrata \S': 9-10.5 mm vs. (9-)12-23 mm; 2: 6-8 mm vs. (6-)16-25 mm(-30) mm;
capsules: 13-19 mm vs. (15-)21-36(-60) mm].

t-UtJey and UtJey: New species a'

Figure 4. Begonia mcphersonii. Holotype (McPherson 12019, MO).

Additional specimens examined. PANAMA. Chiriqui. Above San Felix on mining rd. 18-
27 mi off the Pan Am Hwy above Chame or turnoffto Escopeta, 12 Mar 1976, T. B. Croat 33075
(MO, USF); vie. of Cerro Colorado mine development, 28 mi above San Felix, 9- 10 mi above turnoff
to Escopeta, 13 Mar 1978, Croat 33206 (MO); on rd in vie. of branch in rd. to Cerro Colorado and
Escopeta, above Rio San Felix ca 13 mi N of Rio San Felix bridge, 15 Mar 1976, Croat 33444 (MO).

5. BEGONIA PSEUDOPELTATABurt-Utley & Utley, sp. nov. TYPE. HONDURAS. Lempira.

Trail from old electricity generation plant to Camp Don Tomas ca. "10 straight line" WSW of
Gracias, Parque Nacional Celaque, 14°33'N, 88°40'W, 1850 m, 11 Feb 1993, R Evans 1154
(holotype: MO; isotype: USF). Figure 5.

Suffrutescent herbs with branching stems to 1.5 m tall; upper internodes l-2.9(-5.3) cm, 4-
8 mm diam, sparingly to densely tomentose with uniseriate whiplash trichomes to 4 mm, intermixed
with stout multiseriate whiplash trichomes to 8 mm. Stipules persistent, asymmetrically oblong to
oblong-ovate, 12-14.5 x 7.5-12 mm, apically obtuse to truncate, marginally entire, glabrous to
sparingly tomentose, weakly keeled distally with the keel excurrent apically; petioles (0.6-)2.5-9.5
cm, sparingly to densely tomentose with fine sericeous trichomes to 4 mm; leaf blades oblique,
asymmetric, ovate to lance-ovate, 1.8-11.8 x 0,6-8.1 cm, basally peltate, apically acuminate to
attenuate-acuminate, marginally ciliate-denticulate to ciliate-crenulate and undulate, occasionally
dentate at ends of major nerves; sparingly hirsute above, becoming giabrate, sparingly to moderately
tomentose below with trichomes most dense on major nerves; 7-9-peltinerved. Inflorescences
shorter than to exceeding the leaves, symmetric to asymmetric, densely cymose, ca 40 or more-
flowered; peduncles 3.8-14.3(-22.5) cm, sparingly to moderately tomentose; bracts deciduous, the
lowermost oblong, 4-11 x 2-3 mm, apically ciliate-lacerate, marginally distally ciliate-lacerate,
hirsute Mamissate flowers with pedicels 6-14 mm; sepals 2, obovate to elliptic, 7-10.5 x 3.5-6.5
mm, glabrous; petals 2, narrowly elliptic to obianceolate, (4.5-)7.5-8.5 x 1.6-2.8 mm; stamens 7-13;
filaments 0.8-2 mm, borne on a raised torus; anthers oblong to oblong-obovate, 1.1-1.7 x 0.6-0.9
mm. Pistillate flowers with pedicels (4.5-)9-ll mm; bracteoles wanting; sepals 2, obovate, 7-8 x 4-
5.5 mm, glabrous; petals 1, obianceolate, 5.5-7 x 1.5-2 mm; ovary trilocular with bipartite placentae,
5.5-7.5 mm, glabrous; styles 3, 2-2.5 mm, fused basally; stigmas lunate. Capsules with pedicels 11-
18 mm; bodies 7-9 mm; locules externally appearing broadly elliptic to suborbicular, 4.5-6.5 x 4-7
mm; wings 3, subequal to somewhat unequal, the largest wing weakly asymmetric, broadly triangular
to lunate-triangular, 5-6,5 x 5.5-8 mm, the second and third wings shaliowly lunate to
asymmetrically triangular.

Distribution and habitat. Begonia pseudopeltata is known from Guatemala, Honduras, and
Nicaragua at elevations between 600 and 2000 m, where it lias been collected in mixed pine and
hardwood secondary growth.

Begonia pseudopeltata is an attractive suffrutescent species with abundant small flowers and
capsules. It resembles B. peltata Otto & Dietr. in its peltate leaves and is potentially closely related to
this taxon. Although B. peltata (syn. B. incana Lindl.) was included in sect. Rachia (Kl.) A- DC, this
species was most recently placed in sect. Gireoudia (Doorenbos et al. 1998). Begonia pseudopeltata
is readily distinguished from this latter taxon by its oblong to oblong-ovate stipules (vs. ovate to
triangular) and floral characters, including its more narrow staminate sepals (3.5-6.5 mm vs. 7-11
mm), fewer stamens (7-13 vs. 14-27), longer styles (2-2.5 mm vs. 1.3-2 mm), smaller ovaries (5.5-
7.5 mm vs. 6.5-12 mm), and smaller capsules (7-9 mm vs. 10.5-18 mm). Although these species
share somewhat overlapping distributions, with both occurring in Guatemala and Honduras, B.
pseudopeltata has not been reported from Mexico and B. peltata is not known to occur in Nicaragua.
Moreover, there is no evidence that these species have been collected from the same general localities

III UlllU ' lll.lklll.ll.l '•! 1 1 Illl.

urt-Utley and UUey: New species and notes on Begonia 12

Figure 5. Begonia pseudopeltata. Isotype (Evans 1154, USF).

Additional specimens examined. GUATEMALA. Chiquimula. Cerro Brujo, in vie. of Rio

Negro, belov, Montana Montenegro, near village of Brujo. 1500-2000 m, 1 Nov 1939. Steyermark
30937 (F). HONDURAS. Copan. ca Dulce Nombre, 1200 m, 30 Mar 1963, Molina R. 11 748 (F, LL,
NY). Lempira. Sendero entre La Planta Electrica y las primera casa (abandonada) de Don Tomas
Parque Nac. de Celaque, 14°36'N, 88°40'W, IS Mar 1991, 1800 in. House 892 (MO, USF); Cuyamel,
without further locality. 30 Mar 1923, Carleton 474 (US). NICARAGUA. Jinotega. Las Alturas de
Kilambe, "Filas el Portal" NE del Cerro Kilambe, 13°37'N, 85°40'W, 600-900 m, 26 Mar 1981,
Moreno & Sandino 7594 (MO).

6. BEGONIA AGUABUENENSIS Burt-Utley & Utley, sp. nov. TYPE: COSTARICA. San Jose.

Canton Leon Cortes. Z.P. Caraigres, Cuenca del Pirres-Damas, Cerro Caraigres, Falda SE
Fila Aguabuena, entre Quebrada Pilas y Quebrada Ceniza, 9°42'45'N, 84°06'21"W, 1000-
1200 m, 26 Dec 1996, J. F. Morales 5945 (holotype: MO). Figure 6.

Rhizomatous herbs; internodes short and stout, 0.9-1 cm long, 1.3-1.8 cm diam, squamose

with very broad lacerate and laciniate trichomes 3-5 mm. Stipules persistent, refiexed and revolute
with age, asymmetrically ovate. 13-23 x 9-13 mm, marginally entire, glabrous, strongly keeled with
the keel fimbriate; petioles 8.5-18.5+ cm, glabrous except for a ring of lacerate scales 4-5(-6.5) mm
just beneath the petiole-blade junction; leaf blades ovate, 14-21 x 6.5-18 cm, apically attenuate-
acuminate, basally peltate, marginally ciliate and weakly undulate, but denticulate at ends of major
nerves, above glabrous to minutely glandular, below sparingly squamose on major nerves with
trichomes 1.5-3 mm. 6-7-peltinerved. Inflorescences exceeding the foliage, asymmetrically cymose,
many-flowered; bracts caducous, not seen; peduncles 22-59 cm, glabrous. Staminate flowers with
pedicels 10-12 mm, glabrous; sepals obovate, 5.5-6.5 x 3.5-5.5 mm, glabrous, white-pink; petals
wanting; stamens 7-11, borne on a raised torus; filaments 0.3-0.6 mm; anthers obovate, 1.1-1.3 x
0.5-0.7 mm. Pistillate flowers with pedicels 6-8 mm, glabrous; bracteoles wanting; sq^als 7.5-10 x
5.5-8 mm, glabrous, white-pink; petals wanting; ovaries trilocular with bipartite placentae, 7-8 mm,
glabrous; styles 3, 2-2.5 mm; stigmas lunate. Capsules with pedicels 8-11 mm; bodies 11-14 mm,
locule chambers externally appearing obovate, 9x7 mm; wings subequal, the primary wing
asymmetrically lunate-triangular, 5-7 x 9-11 mm; the other wings lunate, 4.5-6 x 8-9 mm.

Distribution and habitat. Begonia aguabuenensis is known only from central Costa Rica in
San Jose province between 1000-1300 m.

Begonia aguabuenensis (sect. Gireoudia Kl. A. DC.) is distinguished hs its stout rhizomes,
peltate leaf blades, squamose petioles, apetalous staminate and pistillate flowers, and subequal ovary
and capsule wings. It appears most similar to Begonia manicata var peltata L.B. Sm. & B.G. Schub.
from Guatemala. Both taxa share similar rhizornatous habits, leaf blade size, peltate leaf blades,
squamose petiolar trichomes, apetalous flowers, staminate sepals within the size observed for B.
manicata var peltata, ebracteoiate pistillate flowers, and similar capsule size and subequal capsule
wings. The petiolar indument of both taxa is squamose, but that of B, aguabuenensis is restricted to a
band just below ? the blade, while in B. manicata var peltata it is distributed throughout the petioles but
is most dense just beneath the blade. Begonia manicata var peltata is known only from its type and
one additional collection from Guatemala (Burt-Utley 1985). There are no known collections of B.
manicata var peltata from Honduras or Nicaragua, while B. aguabuenensis appears restricted to Costa
Rica. Ideally, more collections of B. manicata var peltata are needed before its synonymy w r ith
Begonia aguabuenensis is made.

Additional specimens examined. COSTARICA, San Jose. Canton de Acosta. Cuenca del
Pirris-Damas, Fila Bustamante, Fila Aguabuena, Aserri, entre Quebrada Chilamate y Quebrada Pilas,
camino, 9°43'35"x84°ll'20"W, 1300 m, 12 Dec 1996, Morales 5921 (MO).

y

m

Figure 6. Begonia aguabuenensis. Holotype (Morales 5945, MO).

7. BEGONIA SUKUTENSIS Burt-Utley & Utley, sp. nov. TYPE: COSTARICA. Limon. Reserva
Indigena Talamanca, Sukut, desembocadura del Rio Sukut en el Rio Uren, camino al SE
hacia Purisqui. 9°24'15"N, 82°58'10 W, 350-550 m, 6 Jul 1989, B. Hammel, LA. Chacon, &
G. Herrera 1 7750 (holotype: MO; isotype: USF). Figure 7.

Herbaceous perennials with stout repent rhizomes; internodes 2-3.8 cm long, 7-1.6 cm
diam, densely tomentose with whiplash trichomes 2-3 mm. Stipules broadly ovate to ovate-
triangular, 18-22 x 14-22+ mm, marginally entire, densely villous or the keel only villous; petioles
25.5-43 cm, minutely glandular and tomentose with whiplash trichomes 1-3 mm; leaf blades oblique,
asymmetrically broadly elliptic to oblong or obovate, 24-32 x 18-24 cm, basally cordate with lobes
occasionally overlapping, apically acuminate, marginally ciliate, finely denticulate, and occasionally
dentate at ends of major nerves; glabrous above except villous above the petiole insertion, villous on
nerves below but only sparingly so in intercostal regions; 14-15-paimatinerved. Inflorescences
exceeding the foliage, asymmetrically cymose, many-flowered; peduncles 1-1.2 m, sparingly pilose
with trichomes 0.3-2 mm and minutely glandular; lowermost floral bracts unknown, but the upper
convex, marginally entire and villous medially. Stammate flowers with pedicels 2-4 mm, villous
and glandular; sepals 2. transversely elliptic, 2-5 x 3-3.5 mm. glabrous to very sparingly villous and
glanduLn cW'Apheies present, white; petals wanting; stamens on a raised torus and occasionally
appearing submonadelphous, 64-75; filaments 0.5-0.7 mm, maroon; anthers broadly obovate to
oblong, 0.3-0.7 x 0.30.4 mm, connective maroon. Pistillate flowers with pedicels 1-4 mm, villous
and glandular; sqjals 2, broadly transversely elliptic to obovate or suborbicular, 4-6 x 3.5-6 mm,
glabrous to sparingly villous and glandular, while, cystospheres present; petals wanting; ovary 2-3
mm, glabrous, cystospheres abundant; styles 3, 0.5-1.2 mm; stigmas bicornute, appearing maroon
when dry. Capsules with pedicels to 2 mm; bodies often beaked, 4.5-5.5 mm; locule chambers
externally appearing suborbicular to broadly ovate, 3-3.5 x 3.5-4 mm; wings 3, unequal, the largest
wing asymmetrically ovate, 7-10 x 5-5.5 mm; the other two wings marginiform, lunate to lunate-
triangular.

Etymology. Begonia sukutensis is named for the region where this species was collected,
Sukut, Reserva Indigena Talamanca, desembocadura del Rio Sukut en el Rio Uren.

Distribution. Begonia sukutensis is known only from the type locality between 350-550 m,
but certainly it is expected elsewhere in Costa Rica and possibly Panama in the appropriate

Begonia sukutensis stands apart from almost all other Mesoamerican species in sect.
Gireoudia (Kl.) A DC. in its very long peduncles, very short pedicels, and very small sepals and
capsules. It is also the only species of Mesoamerican Begonia that has filaments, anthers, stigmas and
styles that appear maroon; in all other species they are yellow (Burt-Utley, pers. obs.). This unique
maroon pigmentation also has not been observed in flowers of western South American taxa (Burt-
Utley, pers. obs.). The only other taxa in the section in Costa Rica and Panama with sometimes small
sepals and submonadelphous stamens like those of B. sukutensis are B. corredorana C. DC, a
suffrutescent species, and the rhizomatous B. quaternata Smith & Schubert (Burt-Utley 1985).
Begonia sukutensis appears most closely related to B. corredorana and occurs at similar elevations,
but the latter species is found in evergreen forests and cloud forests on Costa Rica's and Panama's
Pacific slopes between 20-900 m, while B. sukutensis colonizes Costa Rica's Caribbean slopes. Both
taxa have similar villous-tomentose pubescence but differ in plant habit (rhizomatous vs. caulescent),
stipule form and size, with those of B. sukutensis broadly ovate to ovate-triangular 18-22 x 14-22+
mm (vs. lanceolate, 18-25 x 6-8 mm), glabrous adaxial leaf surfaces (vs. hirsute), more numerous
leaf blade nerves (14 to 15 vs. 10 tol3), elongate peduncles [to 1.2 m vs. (12-)18-36 cm], more
numerous stamens [64-75 vs. 16-41], and its smaller large capsule wings [7-10 x 5-6 mm vs. (12-

)14-19(-22) x (7— )9— 11 mm]. From B, quaternata,B. sukutensis is immediately distinguished by its
tomentose petioles (vs. squamose) and its dichotomously branching inflorescences, in contrast to the
typically 3-6-branched inflorescences characteristic of B. quaternata.

Figure 7. Begonia sukutensis. Isotype (Hammel, Chacon, & He

8. BEGONIA PANAMENSIS Burt-Utley & Utley, sp. nov. TYPE. PANAMA. Chiriqui. Trail

between N fork of Rio Palo Alto and Cairo Pato Macho, 6 km NE of Boquete, 1800-2200 m,
7 Feb 1986, M. Grayum 6418 (holotype: MO; isotypes: PMA, USF). Figure 8.

Herbaceous perennials with slender rhizomes; internodes short to occasionally elongate,
0.7-2 (-4.2) cm, 4-9 (-14) mm diam, glabrous to very sparingly villous with stout trichomes to 1
mm. Stipules caducous to fugacious, ovate to oblong, 11x7 mm, marginally entire, glabrous, only
rarely very sparingly villous, keeled; petioles (4.5— )11— 35 cm, glandular and sparingly pilose at
maturity with fine sericeous villi l-3(-3.5) mm; leaf blades oblique to transversely elliptic or ovate,
(6.5-)9-16.8 x (4-)7.2-ll cm, basally cordate, apically acuminate, marginally eciliate, weakly
undulate, glabrous above and sparingly pilose below, especially on major nerves; 10-12-
palmatinerved. Inflorescences asymmetric, shorter than to exceeding the foliage, ca 8 or fewer-
flowered; peduncles (ll-)16.5-38 cm, sparingly pilose with trichomes 1-3 mm; bracts caducous,
broadly obovate, 17-18 x 20-22 mm, apically refuse, marginally entire, glabrous. Staminate flowers
with pedicels 23-46 mm, glabrous to very sparingly pilose; sepals often coriaceous when dry,
transversely elliptic, 15-21 x 23-30 mm, glabrous to sparingly ^landulai and pilos- piouni'lh
petals wanting; stamens very numerous, in excess of 100; filaments 0,8-3 mm, appearing free or on a
slightly raised torus; anthers narrowly oblong to obovate, (1.3— )1. 8-2.5 x 0.4-0.7 mm. Pistillate
flowers with pedicels 10-15 mm, pilose; bracteoles wanting; sepals transversely elliptic, 10-18 x 17-
27 mm, glabrous to sparingly glandular or very sparingly pilose; petals wanting; ovaries fleshy,
bilocular with bipartite placentae, 11-15 mm, glandular and pilose; styles 3, 4 mm, fused briefly
basally; stigmas bicornute. Capsules with pedicels 19-27 mm; bodies when dry coriaceous before
dehiscence, strongly nutant. weakly to strongly beaked, rarely unbeaked, (15-)17-23 mm; wings 3,
unequal, the largest wing oblong, 14-16 x 15-17 mm; the second and third wings lunate-triangular.

Distribution and habitat. Begonia panamensis o
forests between 1700 and 2300 m. Although all collecti

surprising to find B. panamensis in adjacent western Costa Rica.

Begonia panamensis is a striking rhizomatous species with few flowers and very large sepals.
It has the characteristic fleshy, bilocular ovaries and nutant capsules that dehisce only after their outer
covering has eroded like other species in sect. Weilbachia (Burt-Utley & Utley 1999), a section
known only from Central America and Mexico. Only two other species in this section are known to
occur in Panama, B. carletonii Standi, and B. vestita C. DC. Begonia panamensis is readily
distinguished from these taxa by its apetalous staminate and pistillate flowers, large sepals, and very
numerous stamens. Begonia panamensis is most similar to B. copeyana, a Costa Rican endemic, with
which it shares its apetalous flowers, a similar but less dense indument, and bicornute stigmas.
However, it stands apart from this latter taxon in both vegetative and floral characters, including its
glabrous to very sparingly villous internodes, more numerous primary blade veins (10-12 vs. 7-10),
more numerous stamens [100^ vs. 24-33(-52)], broader staminate sepals (23-30 mm vs. 10.5-20
mm), larger pistillate sepals (10-18 x 17-27 mm vs. 7.5-8 x 10-12 mm), and larger capsules [(15-
)17-23mmvs. 13-16 mm).

Additional specimens examined. PANAMA. Chiriqui. E slopes of Cerro Pando, 8°55'N x
82°44' W, 15 Oct 1981, Knapp 1 666 (USF); 8 km W of Cerro Pando in vicinity of Las Nubes on trail
above stream, 11 Feb 1978, Utley 5669 (DUKE); Cerro Pate de Macho, ca 5 mi NE of Boquete, on
trail to continental divide leading to Finca Serrano, Pacific slopes, 23 Nov 1979, Croat 48559 (MO,
USF); vicinity of Boquete, Cerro Pate de Macho SW slope. 8°46'N, 82°25'W, 19 June 1987, Croat
66418 (MO, USF); trail to top of Cerro Pate de Macho, 8°40'N, 82°28'W, Hoover 557 (MO, USF);
end of rd past Palo Alto to Bocas, Hammel, D 'Arcy, & Averett 6506 (MO); SE slopes and summit of
Cerro Pato Macho, trail from Rio Palo Alto, 4 km NE of Boquete, 27 May 1981, Sytsma, Knapp, &

Burt-Utley and Utley: New species and notes on Begonia \

i, Santa Clara to Cerro Pando, 2S Feb 1985, van der ,

Figure 8. Begonia panamemis. Holotype (from Grayum 6418, MO).

Burt-Utley and Utley: New species and notes on Begonia 19

9. BEGONIA GRACILIOIDES Burt-Utley & Utley, sp. nov. TYPE. GUATEMALA. Solola.

Volcan San Pedro, 2150 m, 20 Sep 1971, A. Molina R. &A.R. Molina 26652 (holotype: F;
isotype: MICH). Figure 9.

Monoecious herbs presumably with underground tubers; leafy stems erect, freely branching,
0.5-1 m tall; often producing clusters of small bulbils 0.4-1.5 mm diam at the nodes; internodes 3.5-
8.5 cm, 3-6.5 mm diam, but the slender branches to 1 mm diam, glabrous to minutely sparingly
glandular. Stipules persistent, asymmetrically broadly ovate, 5.5-8 x 6-9.5 mm, marginally
glandular and ciliate-serrulate, glabrous; petioles (0.6-)2. 1-6.5 cm, hirteilous with trichomes to 1
mm; lower leaf blades oblique to occasionally straight, asymmetrically ovate to deltoid, 4.4-10.3 x 3-
6 cm, basally cordate, apically acute to acuminate, marginally serrate to doubly serrate or dentate,
sparingly ciliate. hirteilous above and below; 6-8-palmatinerved; upper leaf blades asymmetric, much
reduced in size. Inflorescences borne in the axils of upper leaves and terminating the stem,
symmetric, 3-7-flowered; peduncles 2.2-3.8 cm, sparingly minutely glandular; bracts deciduous,
hemiorbicular to obovate, (6-)7-8.5 x 4.5-11 mm, marginally glandular-ciliate-denticulate.
Staminate flowers with pedicels 13 mm, glabrous to minutely glandular; sepals 2, ovate to elliptic,
19-21 x 11-13 mm, apically acute to acuminate, marginally glandular-serrulate to denticulate or
crenulate, glabrous; petals 2, obovate to elliptic, 13 x 8 mm distally glandular-crenulate to denticulate
with short stalked glandular hairs; stamens numerous; filaments to 2 mm, appearing monadelphous;
anthers obovoid to subglobose, inserted at an angle, 0.8-0.9 mm. Pistillate flowers at anthesis
unknown: ebracteolate; tepals presumably 5, elliptic to narrowly ovate, marginally glandular-serrulate
with short stalked glandular hairs, externally glabrous to minutely glandular; ovaries trilocular with
bipartite placentae, 7-10 mm, immediately post-anthesis, glabrous; styles 3, to 1 mm, fused basally;
stigmas bicornute. Capsules with pedicels to 16-18 mm, glabrous to minutely glandular; bodies 17-
18 mm; locules externally elliptic, 14—15 x 7-8 mm; wings 3, unequal, the largest wing
asymmetrically triangular. 11-12 x 16 mm, the second and third wings asymmetrically lunate to
lunate- triangular.

Distribution and habitat. Begonia gracilioides is known only from Guatemala, where it
occurs on moist banks and in thickets between 1800 and 3000 m. This species might actually be
distributed more widely but, because it presumably will die back to its tubers after flowering, will not
be a conspicuous component of the vegetation during part of the year.

Begonia gracilioides (sect. Knesebeckia) is characterized by its branching, leafy erect stems,
bulbils, several-flowered inflorescences, and marginally denticulate to serrulate staminate sepals.
Because bulbils have only been observed on tuberous species from Mexico and Central America, their
presence in leaf axils of B. gracilioides strongly suggests that this species also develops from
underground tubers like the Mexican endemic B. gracilis H.B.K. (pers. obs.). Three other species
known from Central .America with bulbils are also tuberous, B. weberlingii Irmsch. (El Salvador and
Oaxaca, Mexico), B. biserrata Lindl. (Guatemala and Honduras), and B. ignea (Kl.) A DC. (Costa
Rica) (pers. obs.). Although the type and paratype of B. gracilioides were identified previously as B.
gracilis and the species was illustrated in the Flora of Guatemala (Smith & Schubert 1961), this
species differs from B. gracilis in several characters including its several-flowered inflorescences. In
contrast, those of B. gracilis are typically 2-fiow r ered, bearing one staminate and one pistillate flower
(Burt-Utley & McVaugh 200 1). Moreover, peduncles are longer in B. gracilioides than they are in B.
gracilis (2.2-3.8 vs. 0.5-1.5 cm), while inflorescence bracts are deciduous in B. gracilioides but
persistent in B. gracilis. In B. gracilioides, staminate sepals are acute to acuminate and more narrow
than those of B. gracilis (11-13 mm vs. 15-22 mm) and are conspicuously glandular-ciliate and
denticulate, unlike those of B. gracilis, which are simply denticulate to crenulate. Capsules of B.
gracilioides are also smaller than those of B. gracilis (17-18 mm vs. 17-25 mm). These species also
differ in their distributions with B. gracilioides apparently endemic to Guatemala, while B. gracilis is

very widely distributed in Mexico arid has been collected in central Oaxaca, but it is unknown from
Chiapas.

Additional specimens examined. GUATEMALA. Escirintla. Volcan Pacaya, 28 Sep 1972,
Molina & Molina 27652 (F, MICH). Soioia. Volcan Tollman (slopes above San Lucas Tollman), 13
Jun 1 942, Steyermark 47605 (F).

Figure 9. Begonia grac ih 'aides. Holotype (MolinaR. &Molin

Burt-Utley and Utley: New species and notes on Begonia 21

10. BEGONIA TENUIS Burt-Utley & Utley, sp. nov. TYPE. MEXICO. Chiapas. Cerro del
Boqueron, Aug 1913, C.A. Purpus 6937 (holotype: NY). Figure 10.

Herbs presumably developing from underground tubers annually, leafy stems erect to
sprawling, branching weakly distally, 17-40+ cm tall; often producing clusters of small bulbils to 1
mm diam at the nodes; internodes 2.5-7.5 cm, 1^1 mm diam, hirtellous with short villi 0.4-1 mm.
Stipules persistent, ascending to spreading, asymmetrically hemiorbicular to broadly ovate, 2-6 x
3.5-7 mm, marginally ciliate-serrulate to ciliate denticulate, sparingly hirtellous; petioles 1. 1-4.7 cm,
hirtellous with short villous trichomes 0.3-0.5 mm; lower leaf blades asymmetric, obliquely narrowly
ovate, 3.3-11 x 1.5-4.5 cm, basally very shallowly cordate to appearing almost cuneate, apically
attenuate-acuminate, hirtellous throughout above and beneath but trichomes most dense on major
nerves below; 7-8-palmatinerved; upper blades reduced in size. Inflorescences borne in axils of
reduced upper leaves and terminal, 1-2 or more-flowered; peduncles 1.1-3.0 cm, sparingly hirtellous;
bracts obovate to suborbicular, 2-3 x 3-4 mm, marginally ciliate-denticulate to ciliate serrulate,
glabrous to very sparingly hirtellous Staiiiinate flowers with pedicels 10 mm; sepals ovate, 12-14 x
8-10 mm, marginally ciiiate-denticulate to ciliate-serrulate, apparently glabrous; petals 2, elliptic,
12.5-15 x 5.5-7 mm; stamens very numerous, borne on a stout torus, anthers to 1 mm. Pistillate
flowers with pedicels to 14-18 mm; bracteoles wanting, tq^ais 5, variable in shape, elliptic to ovate
or obovate, 7.5-13 x 5-8 mm, marginally the outer 3 ciliate-denticulate to ciliate-serrulate, glabrous;
ovary 8-10 mm, presumably trilocular with bipartite placentae, glabrous; styles 3, 1-1.3 mm fused
briefly basally; stigmas bicornute. Capsules when immature with pedicels to 20 mm; bodies to 1.5
cm; wings 3, apparently subequal.

Etymology. The specific epithet, tenuis, refers to the slender stems of this species.

Distribution and habitat. Begonia tenuis is known only from eastern Chiapas in Mexico,
between 1000-2000 m, but this species should occur in adjacent parts of Guatemala and possibly El
Salvador. Like B. gracilioides, this species would be expected to die back to its tubers after flowering
and therefore may not be evident many months of the year.

Begonia tenuis (sect. Knesebeckia) is characterized by its slender hirtellous stems, bulbils in
its leaf axils, hemiorbicular to broadly ovate stipules and ciliate-denticulate to ciliate-serrulate
staminate sepals. Like B. gracilioides, the presence of bulbils strongly suggests that this species also
develops from underground tubers, since all species that produce bulbils within Mexico and the
region delimited by the Flora Mesoamericana also are tuberous (pers. obs.). In Arthur Houghton's
thesis on the Begoniaceae of North America (Houghton 1924), B. tenuis was described as B. gracilis
var. chiapensis Houghton, but it was never published. This species is most similar to the tuberous
species B. dealbata Liebm. (Oaxaea, Mexico, and Guerrero) in its slender habit, and B. gracilis, also a
Mexican endemic widely distributed through northern, central, and western Mexico. Begonia tenuis
differs from both B. dealbata and B. gracilis in its hemiorbicular to broadly ovate stipules, in contrast
to the unusual basally cuneate and distally lobed, foliaceous stipules characteristic B. dealbata and the
ovate to triangular stipules of B. gracilis. Ovaries of B. tenuis are generally much larger than those of
B. dealbata (8-10 mm vs. 3.5-7 mm) and smaller than those of B. gracilis (8-10 mm vs. 10- 21
mm), while B. tenuis has subequal capsule wings, unlike the unequal capsule wings of both B.
gracilis and B. dealbata (pers. obs.).

Additional specimens examined. MEXICO. Chiapas. Volcan Tacana, Aug 1938, Matuda
6039 (LL, MEXU).

Burt-Utiey and Utiey: New species and notes on Begonia 22

1 f

Figure 10. Begonia tenuis. Holotype (from Purpus 6937., NY).

Burt-Utley and Utley: New species and notes on Begonia 23

Taxonomic Notes

BEGONIA MILITARIS L.B. Sm. & B.G. Schub, Contr. Gray Herb. 154: 24, fig. 2. 1945. TYPE.

GUATEMALA. Alia Verapaz. Chama, 270 m, 15 May 1920, H. Johnson I 78 (holotype:

US!;isotype:F!).
Begonia sciadophora L.B. Sm. & B.G. Schub., Contr. Gray Herb. 161: 28. 1946. TYPE.

GUATEMALA. Alta Verapaz. Dense wet limestone forest near Chirriacte on Peten Hwy ca

900 m, 9 Apr 1941. P. Standley 91967 (holotype: F!).

Begonia militaris is unique among Mesoamerican Begonia in having 5-tepaled pistillate
flowers with bilocular ovaries. Bracteole form, ovary shape, and the species bipartite placentae w ? ere
described, and the illustration in Plate II (Smith & Schubert 1945) is clearly of a nutant capsule;
however, only bilocular ovaries and nutant capsules without bracteoles or bracteole scars were
observed on the specimens examined. These characters together with its creeping rhizomatous habit,
pilose petioles, and peltate leaves easily distinguish B. militaris from all other Mesoamerican Begonia
except B. calderonii Standi. From B. calderonii, B. militaris is readily distinguished by its staminate
flowers with two sepals and two petals (vs. petals wanting) and pistillate flowers with five tepals (vs.
two sepals and no petals). Within Mexico and Central America, other species with bilocular ovaries
and nutant capsules have pistillate flowers with two sepals and 0-1 petals, which is characteristic of
sect. Weilbachia (Burt-Utley & Utley 1999). The affiliation of B. militaris with either a section or
other species remains unclear. Begonia sciadophora is indistinguishable from B. militaris in both
vegetative and floral characters, including its 5-tepaled pistillate flowers with bilocular ovaries and is
synonymized here with B. militaris.

Additional specimens examined. GUATEMALA. Alta Verapaz. Chapultepec Farm, 62
km from Coban on Sebol road, 22 May 1964. Contreras 4763 (DS. US): near Chirriacte on Peten
Hwy, 900 m, 9 Apr 1941, Standley 91953 (F).

BEGONIA LUDICRA A. DC, Ann. Sci Nat. Bot. IV, 11: 133. 1859. LECTOTYPE (designated

here): MEXICO. Veracruz. Cordillera, marais de xalapa, 4000 ft, Jun 1840, Galeotii 189
(G!; isolectotypes: BR!, K!, P!).
Begonia liebmannii A DC, Prodr. 15: 345. 1864. LECTOTYPE (designated here): MEXICO.

Oaxaca. Tonaguia, Aug 1842, Liebmann s.n. (B!; isolectotypes: C[2]!).

Begonia ludicra w r as described by A. DC (1859) without the benefit of capsules and included
in sect. Gireoudia, a section with typically trilocular ovaries. In the Flora of Guatemala (Smith &
Schubert 1961), B. ludicra was also characterized as having "3-cei!ed" ovaries and nutant to very
sharply reflexed capsules. Examination of available collections of B. ludicra establish that the
pistillate flowers have bilocular, nutant, or reflexed ovaries and capsules that must have their outer
coverings erode before dehiscence. These traits are consistent with species in sect. Weilbachia (Burt-
LTtley & Utley 1999) and, with the exception of B. militaris. have not been observed in other taxa in
from Mexico and Central America.

Begonia ludicra has been considered variable in vegetative and floral characters and thought
to be distributed from southern Mexico to Guatemala and Panama (Smith & Schubert 1961).
However, its description and inclusion in the Flora of Guatemala were based primarily on a
vegetatively similar species, B. purpusii Ziesenh. Specimens of this latter taxon consistently were
misidentified as B. ludicra, a species with similar bilocular ovaries and capsules. Begonia ludicra is
readily distinguished from B. purpusii in a number of characters, including its habit and staminate and
pistillate flowers. Begonia ludicra has rhizomes with short to elongate internodes that root at the
nodes and are repent or attached to an upright substrate, where they can form dense masses or mats of

Burt-Utley and Utley: New species and notes on Begonia 24

individuals, while B. purpusii has elongate sterns that may he upright to inclined. Begonia ludicra is
also distinguished from B. purpusii by its staminate flowers with an inner perianth of two petals and
pistillate flowers with one petal. In contrast, staminate and pistillate flowers of B. purpusii have two
sepals and no petals. Begonia ludicra is endemic to Mexico, distributed from the states of Veracruz
and adjacent Puebla to the wet Caribbean slopes of the Sierra Madre in central Oaxaca, while B.
purpusii occurs in eastern Chiapas and Guatemala.

Representative specimens examined. MEXICO. Veracruz. Mpio. Huatusco, Ventura, 3
km NE de Huatusco, 1300 m, 31 May 1979, Avendano & Benavides M. 301 (F); Mpio. Yecuatla,
Lomas de Santa Rila. 3 Jul 1971, Ventura 3621 (MEXU, MICH); near Jalapa. Schiede 733-734 (B);
Jalapa, Jun 1838, Linden 31 (K, MICH). Puebla. Cascada de Oligui, entre Teziutlan y Tlapacoyan,
1550 m, 2 Jun 1968, Garcia Saucedo 76 (MEXU). Oaxaca. Mpio. Sta. Maria Chilchotla, NE de
Agua de Gancho, Agencia Municipal Maria Luisa (8 km del Puente de Fierro, por la terraceria a Sta.
Maria Chilchotla, 18°12'21.6"N, 96°49'28.4" W, 1474 m, 8 Jun 2001, Munn-Estrada & Juarez 1283
(USF); 6 mi S of Puente Nacional on MEX 175 from Tuxtepec to Oaxaca, 2200-2300 ft, 1 Jun 1987,
Utley & Utley 787S (MEXU, USF); 15.1 mi S of Puente at Valle Nacional or 2 mi N of Vista
Hermosa, 4600 ft., 2 Jun 1987, Utley & Utley 7882 (USF); 3.1 mi N of La Esperanza or 16.1 mi S of
bridge at Valle Nacional on Mex 175, 4300 ft, 28 May 1992, Utley & Utley 8777 (MEXU, USF), Dto.
Ixtlan, Puerto San Antonio entre Metates & La Esperanza, 1250 m, 4 Aug 1985 Garcia M., Lorence,
& Allen 1835 (MEXU); 5 km N de Vista Hermosa. km 175 carr. Oaxaca-Tuxtepec, 1260 m, 14 Apr
1982, Torres C. & Lorence 295 (MEXU).

BEGONIA PUSTULATA Liebm., Vid. Meddel. Dansk Naturh. Foren. Kjobenhavn 1852: 6. 1853.
LECTOTYPE (designated here): MEXICO. Oaxaca Lacoba, Distr. Chinantla, Liebmann 202
(C!; isolectotypes: B!, C[2]!).

Begonia pustulata is endemic to Mexico, occurring in eastern Veracruz and the lower

northern slopes of the Sierra Madre in Oaxaca, Its inclusion in the Flora of Guatemala was based on
collections of B. imperialis Lem. that were misidentified as B. pustulata and illustrated there (Smith
& Schubert 1961). This illustration, identified as B. pustulata, was used most recently in a synopsis
of sect. Weilbachia (Doorenbos et al. 1998). Begonia imperialis is more widely distributed than B.
pustulata, ranging from the Isthmus of Tehuantepec to Guatemala between 100 to 1130 m. Although
both species have similar bilocuiar ovaries, the)' are readily distinguished from each other
vegetativeiy and when in flower. Begonia pustulata has larger leaf blades [(7.7-)14-23.3 x (4.6-)8-
16/7 cm vs. 5-14(-16) x 3.5-8(-10.6) cm], generally more nerves [(9-)12-14 vs. (8-)9-10 (-11)],
and longer peduncles [(8.5-)] 5-39 cm vs. (4-)6.2-13.5(-17.3)] than 5. imperialis. Floral characters
also separate these species: B. pustulata lias both staminate and pistillate flowers with two sepals and
an inner perianth of petals, while flowers of B. imperialis lack petals. Although petals occasionally
may abscise from flowers of B. pustulata, all specimens examined and populations visited in Oaxaca
had flowers with an inner perianth series (Burt-Utley, pers. obs.).

Begonia faustinoi Burt-Utley & Utley from Chiapas, Mexico also has been confused with B.
pustulata, with which it shares certain vegetative and floral characters, including a similar villous
indument, staminate and pistillate flowers with both sepals and petals, as well as nutant, bilocuiar
ovaries and capsules. Begonia pustulata can be distinguished from B. faustinoi by its consistently
pustulate or bullate upper leaf surfaces, unlike the even upper surfaces of B. faustinoi, as well as its
more numerous nerves [(9-)ll-14 vs. (8-)9-10(-ll)] and longer peduncles [(8.5-)15-39 vs. (4-)6.2-
13.5(-15.5) cm] (Burt-Utley & Utley 1999).

Representative specimens examined. MEXICO. Veracruz. Mpio. Catemaco, Dos Amantes
entre Catemaco y Sontecomapan, 400 m, 17 Jun 1972, Beaman & Alvarez del Castillo 6199 (F,

Burt-Utley and Utley: New species and notes on Begonia 25

XAL). Oaxaca. 6 mi above Valle Nacional on Hwy 175 to Oaxaca, 23 Aug 1977, Croat 43924
(MO); 17°30'N x 86 ° 30' W, 100 m, 2 Jul 1981, Halm 626 (USF); 5.7 mi S of bridge at Valle
Nacional on Mex 175 from Tuxtq>ec to Oaxaca, 2100 ft, 5 Aug 1987, Utley & Utley 8042 (MEXU,
USF); 5.8 mi S of bridge at Valle Nacional on Mex 175, 2300 ft. 28 May 1992, Utley & Utley 8775
(MEXU, USF); 6 mi above Valle Nacional on Hwy 175 to Oaxaca, 660 m, 23 Aug 1977, Croat 43924
(MO); 6.2 mi S of Valle Nacional on Mex 175, 2400 ft. 30 Dec 1985. Utley & Utley 7525 (USF); 6.2
mi S of Valle Nacional on Mex J 75. 2400 ft, 23 Dec 1986, Utley & Utley 7649 (MEXU, USF); 6.8 mi
S of bridge at Valle Nacional on Mex 175 from Tuxtepec to Oaxaca. 2600 ft, 1 Jun 1987, Utley &
Utley 7873 (MEXU, USF); 12 mi S of bridge at Valle Nacional on Mex 175 from Tuxtepec to
Oaxaca, 100-200 m N of km 68 marker. 3200 ft, 5 Aug 1987. Utley & Utley 8038 (MEXU, USF); 4.5
mi N of Vista Hermosa on Mex 175, 3600 ft, 12 July 1982, Utley & Utley 7046 (USF); near Santiago
Zacatepec, 1500 m, 24 May 1939, Schultes 493 (MEXU).

ACKNOWLEDGEMENTS

Field work was made possible by grants from the Standley Smith Horticultural Trust and the
American Philosophical Society. We are grateful to Dr. Gerrit Davidse for locating additional
Begonia specimens for us, Dr. Carmen Ulloa for providing essential literature, and Dr. Richard
Wunderlin and Dr. Bruce Hansen for their continued support of our research. We thank Mr. Alan
Franck for the digital images of the type specimens, and we especially thank the curators of the
following herbaria for the loan of specimens or use of facilities that made this research possible: B,
BM BR, C, CAS, CR F, GH, K, LL, MEXU, MICH, MO, NY, R US, USE and XAL,

LITERATURE CITED

Burt-Utley, K. and J.F. Utley. 2011. New species and notes on Begonia (Begoniaceae) from Middle

America, I. Novon 21:393^101.
Burt-Utley, K. and J.F. Utley. 1999. Contributions toward a revision of Begonia section Weilbachia

(Begoniaceae). Novon 9: 483^189.
Burt-Utley, K. and R McVaugh. 2001. Begoniaceae. In R McVaugh and W.R Anderson (eds.),

Flora Novo- Galiciana 3: 653-695.
Burt-Utley, K. 1985. A revision of the Central American species of Begonia section Gireoudia

(Begoniaceae). Tulane Stud. Zool. andBot. 25: 1-131.
Doorenbos, J., MS.M Sosef, and J.J.F.E. de Wilde. 1998. The sections of Begonia including

descriptions, keys and species lists. (Studies in Beeoniaceae VI). Wageningen Agric. Univ.

Pap. 98(2): 1-266.
Houghton, AD. 1924. A Monograph of the Begoniaceae of North America. Ph.D. thesis. Univ. of

California, Berkeley.
Smith, L.B. and B.G Schubert. 1946. Begoniaceae. Flora of Colombia. Caldasia 4, 16: 3-38.
Smith, L.B. and B.G. Schubert. 1961. Begoniaceae. Flora of Guatemala. Fieldiana Bot. 24: 157-185.