Botanical Studies (2006) 47: 97-110.

*

Corresponding author: E-mail: bopeng@sinica.edu.tw; Tel:

+886-2-2789-3821; Fax: +886-2-2789-1623.

Three new taxa of

Bego nia

(sect.

Co elo centru m

,

Begoniaceae) from limestone areas in Guangxi, China

Ding FANG

1

, Shin-Ming KU

2, 3

, Yi-Gang WEI

4

, De-Hai QIN

1

, and Ching-I PENG

2,*

1

Guangxi Institute of Traditional Medical and Pharmaceutical Sciences, Guangxi Zhuangzu Autonomous Region,

Nanning 530022, China

2

Herbarium, Research Center for Biodiversity, Academia Sinica, Nangang, Taipei 115, Taiwan

3

Department of Life Sciences, National Cheng-Kung University, Tainan 701, Taiwan

4

Guangxi Institute of Botany, Guangxi Zhuangzu Autonomous Region and Academia Sinica, Guilin 541006, China

(Received July 18, 2005; Accepted December 3, 2005)

ABSTRACT.

Begonia ningmingensis, B. ningmingensis var. bella and B. retinervia, three new taxa of B.

sect. Coelocentrum from the karst area in Guangxi Zhuangzu Autonomous Region, China, are described and

illustrated. Notably, their leaves are adorned with white or pale maculation along all major veins. Begonia

ningmingensis var. ningmingensis resembles B. morsei Irmsch., differing by having larger leaves with white

maculation on the major veins and having larger staminate flowers. The maculation pattern in the leaves of

B. morsei is sharply distinct: they are adorned with a broad, white ring between the center of the leaf and

the margin. Begonia ningmingensis var. bella differs from the typical variety mainly by the suborbicular or

reniform leaves with a rounded or obtuse apex (vs. broadly ovate or suborbicular with an acuminate, shortly

acuminate or acute apex). Begonia retinervia is distinct chiefly by having thickly chartaceous leaves, the

broadly trigonous-ovoid or obovoid ovary and fruit covered with crisp-villous hairs, and the ventral surface

markedly convex, and in the much protruded abaxial wing.

Keywords: Begonia morsei; Begonia ningmingensis; Begonia ningmingensis var. bella; Begonia retinervia;

Begoniaceae; Begonia sect. Coelocentrum; China; Chromosome number; Guangxi; Limestone flora; New

species; New variety.

INTRODUCTION

Begonia sect. Coelocentrum comprises about 18 species

that are distributed mainly in southern China, but also in

northern Vietnam (Shui et al, 2002). During the botanical

inventories undertaken in recent years, many new species

were discovered in China: Ku et al. (2004: B. curvicarpa

and B. luochengensis); Liu et al. (2005: B. picturata); Peng

et al. (2005a: B. fangii; 2005b: B. liuyanii), and Nguyen

(2004) published B. phuthoensis from Vietnam. More

recently, Shui and Chen (2005) added 11 new species and

three new varieties for B. sect. Coelocentrum in China and

proposed a new name, B. variegata for B. masoniana var.

maculata S. K. Chen et al. from Vietnam. Begonia sect.

Coelocentrum is therefore much more speciose than was

previously believed. In this study we report two additional

new species and an additional new variety, all with showy

leaves, from the limestone areas in Guangxi, China.

New taxa

1.

Begonia ningmingensis D. Fang, Y. G. Wei & C.-I

Peng, sp. nov. ¡XTYPE: CHINA. Guangxi Zhuangzu

Autonomous Region, Ningming Xian, Tingliang

Xiang, Longrui Nature Protected Area, elev. ca. 140

m, broad-leaved forest, on limestone rock face in

creek, shaded, wet, occasional, living collection, 2

Sep 2002, type specimens (in flower and fruit) pressed

from plants cultivated in experimental greenhouse

at Academia Sinica, 18 Oct 2004, Wai-Chao Leong,

Yan Liu & Shin-Ming Ku 3410-A (holotype: HAST;

isotypes: A, GXMI, IBK, MO, PE). ¹ç©ú¬î®ü´Å

Figures 1, 2

Begonia morsei auct. non Irmsch.: Ku, T. C., Fl. Reipubl.

Popularis Sin. 52(1): 139, pl. 30: 3, 4. 1999, excl. typ.

Begoniae morsei similis, sed a qua differt praecipue

foliis majoribus, decoratis cum macula alba secundum

venas majores, et floribus staminatis majoribus.

Herbs monoecious; epipetric; perennial; rhizomatous.

Rhizome 2-10 cm or more long, (2.5-)5-10(-16) mm

thick, internodes (3-)5-10(-17) mm long, brown, reddish

brown or greenish brown, villous near petiole bases and

TAXONOMY

98

Botanical Studies, Vol. 47, 2006

Figure 1. Begonia ningmingensis D. Fang, Y. G. Wei & C.-I Peng var. ningmingensis. A, Habit, leaf to right of inflorescence

showing vestiture, leaves to left showing maculation. B, Leaf margin, adaxial surface; B', abaxial surface. C, Portion of leaf, adaxial

surface; C', abaxial surface. D, D', Stipules. E, Bracts. F, Staminate flower. G, Stamen, lateral view; G', abaxial view; G", adaxial

view. H, Carpellate flower, adaxial view; H', lateral view. I, Style and stigma, adaxial view; I', lateral view. J, Capsule. K: a-g, Serial

cross sections of immature capsule. L. Seed. [D, L from Peng et al. 20320, HAST; others from Leong et al. 3410, HAST. All drawn

from plants cultivated in the experimental greenhouse at Academia Sinica.]

FANG et al. ¡X Three new taxa in

Begonia

sect.

Coelocentrum

in China

99

Figure 2. Begonia ningmingensis D. Fang, Y. G. Wei & C.-I Peng var. ningmingensis. A, B, C, Habit and habitat. D, Cultivated

plant at anthesis. E, Stipule. F, Leaf, cross section. G, Staminate flower, lateral view. H, Staminate flower, adaxial view. I, Carpellate

fower, lateral view. J, Carpellate fower, adaxial view. K, Middle cross section of immature capsule. L, Style. M. Capsule. N, Seed

SEM microphotograph. [A from L i HZ-047 (KUN); B from Leong et al. 3632 (HAST); C, N from Peng et al. 20320 (HAST); D-M

from greenhouse-grown plants of Leong et al. 3410-A (type collection, HAST).]

100

Botanical Studies, Vol. 47, 2006

leaf scars. Stipules eventually deciduous, triangular,

narrowly triangular or ovate-triangular, (4-)5-10(-16)

mm long, 4-10 mm wide, greenish or reddish hyaline,

herbaceous or membranaceous, weakly keeled, with or

without reddish nerves, with sparse minute subsessile

brown glandular hairs on abaxial surface, midrib and base

sparsely villous or glabrous, margin serrulate and ciliate

or ciliolate, apex aristate, arista

0.5-3 mm long, hair-

like. Leaves 2-8, alternate, simple, asymmetric, unlobed

or rarely very shallowly angular- lobed, broadly ovate

or suborbicular, base cordate, margin denticulate and

ciliolate, apex acuminate, shortly acuminate, or acute,

rarely obtuse, (5.5-)10-15(-19) cm long (basal lobes

included), (5-)7-12(-13) cm wide, upper surface green,

dark green or dark brown with white maculation (these

areas pale green abaxially) along major veins, sometimes

only with some small white dots along major veins, very

rarely without maculation, lower surface usually reddish,

texture papery or thinly so, surface rugose or rugulose,

adaxially pilose-setose or villous-setose (trichomes 1-1.5

mm long, whitish hyaline or reddish, with a swollen base),

abaxially shortly villous, particularly pronounced on

veins;

venation basally (6-)7(-8)-palmate, midrib distinct,

veins pinnate along midrib, with 1-3 major lateral veins on

each side, other primary veins branching dichotomously

or nearly so, tertiary veins loosely reticulate and weakly

percurrent, forming a divergence angle of 60-85¢X to major

veins, minor veins reticulate, all veins on abaxial surface

prominently raised;

petiole terete,

(3-)8-15(-24) cm long,

2-4(-5) mm thick, brownish or brownish red, villous.

Inflorescences axillary, 1-6 or more, arising directly from

rhizome, dichasial cymes branched 2 or 3(or 4) times,

with 5-21 flowers; staminate flowers 3-17, carpellate

flowers 2-4; peduncle well developed, terete, erect

or

ascending, 4-21 cm long, 1.5-2 mm thick, brownish,

reddish brown or greenish, glabrous, subglabrous or

very sparsely villous.

Bracts caducous, ovate, oblong or

ligulate,

margin denticulate and ciliate or ciliolate (with a

minute, glandular tip), apex obtuse or acute,

2-7 mm long,

1.5-3.5 mm wide, greenish or brownish, membranous.

Staminate flowers: pedicel 7-20 mm long, glabrous or with

few minute subsessile glandular hairs, erect or ascending;

tepals 4, outer 2 ovate, obovate, oblong or suborbicular,

base rounded or slightly cordate, margin entire, eciliate,

apex rounded,

12-20(-25) mm long,

12-16(-19) mm wide,

pinkish, white or greenish white, with very sparse minute,

subsessile, brown glandular hairs on outer surface, white

or pinkish on inner surface; inner 2 tepals oblanceolate or

narrowly obovate (rarely obovate), base cuneate, margin

entire, eciliate, apex obtuse, 8-12(-18) mm long, 3-5

mm wide, white, glabrous; androecium zygomorphic,

subglobose,

stamens (25-)40-50, golf-club-shaped;

filaments subequal, 1.5-3 mm

long, nearly free; anthers ¡Ó

upwardly facing, locules 2, slightly compressed, oblong-

obovoid, apex of connective slightly emarginate, 1.2-1.7

mm long,

0.7-0.8 mm wide, yellow. Carpellate flowers:

pedicel 5-12 mm long, horizontally spreading to pendent,

bracteole absent or sometimes with 1 tiny bracteole; tepals

3, caducous, if persistent not thickened when fruiting,

outer 2 tepals suborbicular or broadly obovate, margin

entire, eciliate, 10-14 mm long, 8-14 mm wide, white or

pinkish, with sparse, minute, subsessile, brown glandular

hairs on outer surface; inner tepal oblanceolate, obovate or

narrowly so, base cuneate, apex obtuse or acute,

7-8 mm

long,

2.5-4 mm wide, white, glabrous; ovary trigonous-

ellipsoid,

7-10 mm long, 4-5 mm thick (wings excluded),

greenish white or reddish white, with or without very few

glandular pilose trichomes (0.2-0.6 mm long) and sparse,

minute, subsessile, brown glandular hairs, 3-winged;

wings unequal or subequal; lateral wings 1.5-2 mm tall;

abaxial wing crescent-shaped or nearly so, 2-4 mm tall,

6-9 mm wide; locule 1, placentation intruded parietal

(axile at base); placentae 3, each 2-branched; styles 3,

fused in lower 1/4-1/2, yellow, ca. 2 mm long,

apically

C-shaped; stigmatic band slightly spiraled. Capsule

nodding, stipe 6-20 mm long, trigonous-ellipsoid, 9-10

mm long, 4-6 mm thick (wings excluded), greenish or

reddish when fresh, apex with persistent styles; lateral

wings 2-3.5 mm tall; abaxial wing crescent-shaped or

nearly so,

4-6 mm tall.

Seeds numerous, brown, ellipsoid

or broadly so,

0.4-0.5 mm long, 0.28-0.32 mm thick,

chalazal end rounded, micropylar end obtuse or slightly

constricted,

outer periclinal walls of mature seeds concave;

collar cells elongated, straight, nearly rectangular, 11-13

cells in a ring, occupying 1/4-2/3 of seed length. Somatic

chromosome number, 2n = 30 (Figure 3: A).

Additional specimens examined. CHINA. Guangxi

Zhuangzu Autonomous Region: Ningming Xian, Tingliang

Xiang, Yuanjing Village, in dense broadleaved forests

on limestone hill slope, elev. ca. 350 m, 10 Nov 1979,

Nonggang Compl. Exped. 20849 (GXMI); Ningming

Xian, Tingliang Xiang, Longrui Nature Protected Area,

elev. ca. 140 m, broadleaved forest, on limestone rock

face in creek, shaded, wet, occasional, 2 Sep 2002, W. C.

Leong, Y. Liu & S. M. Ku 3410 (HAST); Ningming Xian,

Tingliang Xiang, Yuanjing Village, limestone hills, 30 Oct

2002, Y.-G. Wei 0223; Longzhou Xian (Longjin formerly),

Paizong Xiang, Longxu, in dense forests on a limestone

hill top, S. Q. Chen 13915 (IBK, IBSC); Longzhou Xian,

Nonggang National Nature Reserve Headquarters, in a

shaded, moist environment, 9 Nov 1985, Z. Y. Li, H. N.

Qin & D. Z. Fu 005 (PE); Longzhou Xian, Shanglong

Xiang, Nonggang National Nature Reserve, 22¢X29¡¦20¡¦¡¦

N, 106¢X57¡¦29¡¦¡¦E, elev. ca. 180 m, mixed bamboo and

broad-leaved forest, on limestone rock face, shaded,

moist, occasional, 31 Aug 2002, W. C. Leong, Y. Liu &

S. M. Ku 3404 (HAST); Longzhou Xian, Jinlong Zhen,

Nonggang National Nature Reserve, 22¢X27¡¦33¡¦¡¦N, 106¢X

58¡¦01¡¦¡¦E, broadleaved forest, in small valley, on limestone

rock, shaded, moist, rare, 13 Sep 2003, W. C. Leong, J.

Li, X. W. Li, W. H. Chen & Y. M. Shui 3632 (HAST);

Longzhou Xian, Xiangshui Zhen, 3 Jun 2002, Li HZ-047

(KUN); Chongzuo Xian, Laituan Zhen, Longfeng Cun,

White-headed Langur (Trachypithecus leucocephalus)

Reserve, in sparse forests on limestone hill slope, elev.

ca. 170 m, 23 Aug 2004, White-headed Langur Reserve

FANG et al. ¡X Three new taxa in

Begonia

sect.

Coelocentrum

in China

101

Figure 3. Mitotic chromosome spread of Begonia. A, Begonia ningmingensis var. ningmingensis (2n = 30); B, Begonia ningmingensis

var. bella (2n = 30); C, Begonia retinervia (2n = 30).[A from W. C. Leong et al. 3410, B from C.-I Peng et al. 19763, C from C.-I

Peng et al. 19820. Vouchers at HAST.]

Compl. Exped. B0445 (IBK); Chongzuo Shi, Laituan

Zhen, Longfeng Cun, Dingna Tun, 22¢X24¡¦20¡¦¡¦N, 107¢X

30¡¦38¡¦¡¦E, elev. ca. 175 m, on limestone rock face, N- or

E-facing, occasional, 12 Mar 2005, C.-I Peng, S. M. Ku

& H. Y. Chen 20320 (HAST); Chongzuo Shi, Laituan

Zhen, Longfeng Cun, Dingna Tun, 22¢X24¡¦17¡¦¡¦N, 107¢X30¡¦

11¡¦¡¦E, elev. ca. 175 m, on limestone rock face, shaded or

semishaded, moist to dry, occasional, 12 Mar 2005, C.-I

Peng, S. M. Ku, H. Y. Chen 20322 (HAST)..

Ecology. On limestone hills in broadleaved forests.

Distribution. Southwestern Guangxi, China (Figure 4).

Etymology. The specific epithet is derived from the

type locality, Ningming Xian (Ningming County).

Phenology. Flowering from August to November;

fruiting from October to January.

Notes. Upon checking the specimen [Z. Y. Li et al. 005

(PE 1370622)] on which the line drawing of "Begonia

morsei Irmsch." in Ku (1999, pl. 30: 3, 4) was made, we

determined that it is identifiable to B. ningmingensis var.

ningmingensis. The expression of the leaf maculation of

B. ningmingesis var. ningmingensis is variable (see Figure

2: A, B, C, D) and the leaves of plants in cultivation are

often larger than those in the field. The maculation of the

leaves may also be more pronounced in cultivated plants.

Begonia ningmingensis var. ningmingensis resembles B.

morsei Irmsch., differing mainly by having larger leaves

with white maculation along the major veins and having

larger staminate flowers. The maculation pattern of the

leaves of B. morsei is sharply distinct: it consists of a

broad white ring between the center of the leaf and the leaf

margin (see Liu et al., 2005: Figure 7). Begonia morsei is

known from only two specimens (syntypes) collected by

H. B. Morse, without indication of the collection dates.

Begonia morsei has not been recollected since the name

was published by Irmscher (1939). Table 1 summarizes

salient morphological features of B. ningmingensis var.

ningmingensis, B. ningmingensis var. bella, B. retinervia

and B. morsei.

2.

Begonia ningmingensis var. bella D. Fang, Y. G.

Wei & C.-I Peng, var. nov. ¡XTYPE: China. Guangxi

Zhuangzu Autonomous Region, Daxin Xian, Leiping

Zhen, on shady limestone hills, 28 Oct 2002, Yi-Gang

Wei 0221 (holotype: GXMI; isotypes: GXMI, IBK,

PE).

ÄR¸­¬î®ü´Å

Figures 5, 6

Differt a varietate typica Begoniae ningmingensis foliis

suborbicularibus vel reniformibus apice rotundatis vel

obtusis.

Herbs monoecious; epipetric; perennial; rhizomatous.

Rhizome 2-10 cm long, 3-5(-10) mm

thick,

internodes 3-7

mm long, brown or reddish brown,

villous near petiole

bases and leaf scars. Stipules

eventually deciduous,

triangular or ovate-triangular,

5-8 mm long, 2.5-7

Figure 4. Distribution of Begonia ningmingensis var.

ningmingensis (Ė ), B. ningmingensis var. bella (

„\

) an d B.

retinervia („I ) in Guan gxi Z huangzu Autono mous Regi on,

China.

102

Botanical Studies, Vol. 47, 2006

Figure 5. Begonia ningmingensis var. bella D. Fang, Y. G. Wei & C.-I Peng. A, Habit, upper two leaves showing maculation and

others showing vestiture. B, Leaf, showing venation (left half), indumentum (upper right) and maculation (lower right). C, Portion

of leaf, adaxial surface; C', cross section; C", abaxial surface. D, Stipule, abaxial surface; D', adaxial surface. E, Bract. F, Staminate

flower. G, Androecium, showing filaments partly fused at base; G', showing nearly free filaments. H, Stamen, abaxial view; H', side

view; H", adaxial view. I, Carpellate flower, adaxial view; I', lateral views. J, Style and stigmatic band, abaxial view; J', adaxial view. K:

a-e, Serial cross sections of ovary. L, Capsule. M, Seeds. [All from Peng et al. 19763 (HAST); B, M drawn from field material; others

from plants cultivated in the experimental greenhouse at Academia Sinica.]

FANG et al. ¡X Three new taxa in

Begonia

sect.

Coelocentrum

in China

103

Figure 6. Begonia ningmingensis var. bella D. Fang, Y. G. Wei & C.-I Peng. A, B, Habit and habitat. C, Cultivated plant at anthesis.

D, Leaf. E, Stipule. F, Staminate flower, outer tepal margin eciliate. G, Staminate flower, outer tepal margin ciliate. H, Carpellate

flower. I, Middle cross section of ovary. J, Capsule, with glandular pilose trichomes. K, Nearly glabrous capsule. L, Seed SEM

microphotograph. [All from Peng et al. 19763 (HAS T); A, B, K, L from field material; others from living plants cultivated in the

experimental greenhouse at Academia Sinica.]

104

Botanical Studies, Vol. 47, 2006

mm wide, greenish or reddish hyaline, herbaceous

or

membranaceous, keeled or weakly keeled, abaxially

with

sparse minute subsessile brown glandular hairs and

sparsely villous or not

along keel,

margin entire, eciliate,

apex aristate, arista 0.5-2 mm long, hair-like. Leaves

2-8, alternate, simple, asymmetric or pseudosymmetric,

unlobed,

suborbicular or reniform, base cordate,

margin

denticulate and ciliolate,

apex rounded or obtuse,

(2.8-)6-8.5(-13.5) cm long (basal lobes included),

(2.5-)5-7.5(-12.5) cm wide,

upper surface dark green,

brown or dark brown, with white maculation (these areas

pale green abaxially)

along major veins,

rarely only

with some small white dots along major veins, lower

surface reddish or red, texture papery, surface rugulose

or rarely bullate,

adaxially villous-setose or setose-

tomentose (trichomes 0.7-1 mm long, whitish hyaline or

reddish, with a swollen base), abaxially shortly villous,

particularly on veins; venation basally 6-8-palmate, midrib

distinct or sometimes indistinct, major veins branching

dichotomously or nearly so,

tertiary veins loosely

reticulate and weakly percurrent, forming a divergence

angle of 60-85¢X to major veins, minor veins reticulate,

all veins on abaxial surface prominently raised; petiole

terete, (2-)6-10(-14) cm long, 2-3(-5) mm thick, brownish

or brownish red, villous. Inflorescences axillary, 1-3,

arising directly from the rhizome, dichasial cymes 1-3(-4)

times branched, flowers 3-15; staminate flowers 2-10,

carpellate flowers 1-4;

peduncle well developed, terete,

erect,

4-15 cm long,

1.5-2 mm thick, brownish, reddish

brown or greenish, glabrous or subglabrous. Bracts

caducous, broadly ovate, obovate or oblong, margin

denticulate and ciliate or ciliolate (with a minute glandular

tip), apex obtuse or acute, 2-5.5 mm long, 1.5-4 mm

wide, greenish or brownish, membranaceous. Staminate

flowers: pedicel 1-2 cm long, glabrous or rarely with few

minute subsessile glandular hairs, erect or ascending;

tepals 4, outer 2 suborbicular to oblate, base rounded or

slightly cordate, margin eciliate or ciliate, apex rounded,

8-13 mm long and wide, abaxially white, pinkish white

or greenish white, with sparse minute subsessile brown

glandular hairs, adaxially white or slightly pinkish, margin

sometimes greenish; inner 2 tepals obovate or narrowly

obovate, base cuneate, margin entire, eciliate, apex

obtuse or rounded, 7-8 mm long, 2.5-3.5 mm wide, white,

glabrous; androecium zygomorphic, subglobose, stamens

20-35, golf-club-shaped; filaments subequal, 1-1.5 mm

long, nearly free or partly fused; anthers locules 2, slightly

compressed, oblong-obovoid, connective apex slightly

emarginate, 1-1.2 mm long, 0.6-0.7 mm wide, yellow.

Carpellate flowers: pedicel 5-12 mm long, glabrous or

with few minute subsessile glandular hairs, horizontal

to pendent, bracteole absent; tepals 3, persistent; outer

2 tepals oblate or suborbicular, margin entire, eciliate

or occasionally ciliolate below middle, 6-10 mm long,

8-10.5 mm wide, abaxially white, pinkish or greenish,

with sparse minute subsessile brown glandular hairs; inner

tepal obovate or narrowly so, base cuneate, apex obtuse or

rounded, 4-7 mm long, 1.5-3.5 mm wide, white, glabrous;

ovary trigonous-ellipsoid, 4.5-7 mm long, 2-5 mm thick

(wings excluded), greenish or reddish, glandular pilose or

not (trichomes 0.2-0.6 mm long) and with sparse minute

subsessile brown glandular hairs; wings 3, unequal or

subequal, lateral wings narrower, 2-3 mm tall, abaxial

wing crescent shaped or nearly so, 3-5 mm tall, 4-6 mm

wide; locule 1, placentation intruded parietal (axile at

base), placentae 3, each 2-branched; styles 3, fused in

lower 1/4 or nearly free, yellow, 2-2.5 mm long, apically

C-shaped, stigmatic band slightly twisted. Capsule

nodding, stipe 6-12 mm long, greenish or reddish when

fresh, 7-9 mm long, 4-6 mm thick (wings excluded), apex

with persistent tepals and styles; lateral wings 2-3.5 mm

tall; abaxial wing crescent shaped or nearly so, 3.5-5

mm tall. Seeds numerous, brown, ellipsoid or broadly

so, 0.43-0.5 mm long, 0.24-0.35 mm thick, chalazal end

rounded, micropylar end obtuse or slightly constricted,

outer periclinal walls of mature seeds concave; collar

cells elongated, straight, nearly rectangular, 13-17 cells

in a ring, occupying 1/3-2/3 of seed length. Somatic

chromosome number, 2n = 30 (Figure 3: B).

Additional specimens examined. CHINA. Guangxi

Zhuangzu Autonomous Region: Daxin Xian, Leiping

Zhen, ca. 250 m, in sparse forests on limestone hill slope,

rare, D. Fang & L. Zeng 0415 (GXMI); Daxin Xian,

Leiping Zhen, 22¢X38¡¦39¡¦¡¦N, 107¢X05¡¦47¡¦¡¦E, ca. 230 m,

on W-facing limestone rocky slope, exposed, dry, 27

May 2005, C.-I Peng, W. C. Leong, S. M. Ku & Y. Liu

19763 (HAST); same field data but specimens (in flower

and fruit) pressed from plants cultivated in experimental

greenhouse at Academia Sinica, 6 Sep 2005, C.-I Peng, W.

C. Leong, S. M. Ku & Y. Liu 19763-A (HAST).

Ecology. On limestone hills, ca. 230-250 m altitude.

Distribution. Southwestern Guangxi, China; very rare

(Figure 4).

Etymology. The epithet "bella" depicts the beautiful

leaves with white maculation along the major veins on the

upper surface with contrasting red lower surface.

Phenology. Flowering from September to October;

fruiting from November to December.

Notes. Begonia ningmingensis var. bella is known

only from one locality in southwestern Guangxi. We

decided to recognize it as a variety of B. ningmingensis

partly because it occurs near the distribution range of var.

ningmingensis and partly because the two entities differ

mainly by leaf shape and size (Table 1).

Key to the varieties of B. ningmingensis

1a.

Leaf broadly ovate or suborbicular with an acuminate,

shortly acuminate or acute apex; stipules serrulate

and ciliate or ciliolate at margin; tepals of carpellate

flowers caducous, if persistent not thickened when

fruiting.................B. ningmingensis var. ningmingensis

1b.

Leaf suborbicular or reniform with a rounded or obtuse

apex; stipules entire, eciliate; tepals persistent and

thickened in fruit................B. ningmingensis var. bella

FANG et al. ¡X Three new taxa in

Begonia

sect.

Coelocentrum

in China

105

Table 1. Comparison of Begonia ningmingensis var. ningmingensis, B. ningmingensis var. bella, B. retinervia and B.

morsei.

B. ningmingensis var.

ningmingensis (Figures

1, 2)

B. ningmingensis var.

bella (Figures 5, 6)

B. retinervia (Figures 7,

8)

B. morsei (Liu et al., 2005:

Figure 7)

Stipules

Margin serrulate and

ciliate or ciliolate;

weakly keeled

Margin entire, eciliate;

keeled or weakly

keeled

Margin entire, eciliate;

keeled or weakly keeled

Margin nearly entire,

eciliate; distinctly keeled

Leaf

Shape

Broadly ovate or

suborbicular

Suborbicular or reniform Suborbicular

Broadly ovate to

suborbicular

Apex

Acuminate, shortly

acuminate or acute,

rarely obtuse

Rounded, rarely obtuse Obtuse, rounded, acute or

shortly acuminate

Shortly acuminate

Size (cm)

(5.5-)10-15(-19) ¡Ñ (5-)

7-12(-13)

(2.8-)6-8.5(-13.5) ¡Ñ

(2.5-) 5-7.5(-12.5)

(4-)8-14(-25) ¡Ñ (3.5-)

7-13(-22)

5-9 ¡Ñ 4-7

Maculation pattern With white bands along

major veins

With white bands along

major veins

With white bands along

major veins

With a broad white ring

between leaf center and

margin

Texture

Papery or thinly so Papery

Thickly papery

Thinly papery

Peduncle

Glabrous or with a few

pilose hairs

Glabrous or with a few

pilose hairs

Glabrous or very sparsely

villous

Glabrous

Tepals

Size in staminate

flowers (mm)

Outer two 12-20(-25) ¡Ñ

12-16(-19); inner two

8-12(-18) ¡Ñ 3-5

Outer two 8-13 ¡Ñ 8-13;

inner two 7-8 ¡Ñ 2.5-3.5

Outer two 6-16 ¡Ñ 7-15;

inner two 6-14 ¡Ñ 2.5-4

Outer two 6-10.5 ¡Ñ 6.5-10;

inner two 4.5-7 ¡Ñ 2-3

Size in carpellate

flowers (mm)

Outer two 10-14 ¡Ñ

8-14; inner one 7-8 ¡Ñ

2.5-4

Outer two 6-10 ¡Ñ

8-10.5); inner one 4-7

¡Ñ 1.5-3.5

Outer two 7-11 ¡Ñ 9-13;

inner one 5-10 ¡Ñ 2.5-4

Not available

Color

White or pinkish

White, pinkish white or

greenish white

Pink to pinkish white Unknown

Persistence in

arpellate flowers

Caducous or persistent

and not thickened when

fruiting

Persistent and slightly

thickened when

fruiting

Persistent and slightly

thickened when fruiting

Unknown

Androecium

Subglobose; anthers

more or less upwardly

facing

Subglobose; anthers

slightly upwardly

facing

More or less compressed;

all anthers pointing

in same direction and

upwardly facing

Subglobose; anthers

slightly upwardly facing

Filaments

Nearly free

Nearly free, occasionally

fused at base into a

short androphore (ca.

0.6 mm)

Nearly free

Fused at base into an

androphore 0.7-1.25 mm

Styles

Fused in lower 1/3

-

1/2,

rarely nearly free

Fused in lower 1/4 or

nearly free

Nearly free

Unknown

Capsule

Shape

Trigonous-ellipsoid,

beak absent or

indistinct

Trigonous-ellipsoid,

beak absent or

indistinct

Broadly trigonous-ovoid

or obovoid, adaxial side

distinctly convex

,

apex

sometimes with a short

beak

Trigonous-ellipsoid, beak

absent or indistinct

Indumentum

Glabrous or sparsely

pilose and with minute

subsessile glandular

hairs

Glabrous or pilose and

with minute subsessile

glandular hairs

Crisp villous and with

minute subsessile

glandular hairs

Only with minute

subsessile glandular hairs

Abaxial wing shape Crescent-shaped or

nearly so, not curved

Crescent-shaped or

nearly so; not curved

Obliquely triangular or

ligulate, rarely obovate-

oblong; ¡Ó curved or

subcucullate

Crescent-shaped or nearly

so; not curved

Height

4-6 mm

3.5-5 mm

(7-)10-15 mm

3-4 mm

106

Botanical Studies, Vol. 47, 2006

3.

Begonia retinervia D. Fang, D. H. Qin & C.-I

Peng, sp. nov.¡XTYPE: CHINA. Guangxi Zhuangzu

Autonomous Region, Du¡¦an Xian, Xiao Xiang, in

sparse forests, on limestone hill slope, elev. ca. 200

m, 30 Sep 2004, Ding Fang & De-Hai Qin 29620

(holotype: GXMI). ¬ð¯ß¬î®ü´Å Figures 7, 8

Begoniae ningmingensis similis, sed a qua differt

praecipue foliis crasse chartaceis, ovario et fructu late

trigono-ovoideo vel obovoideo et crispo-villoso, et ala

abaxiali multum protrudenti.

Herbs monoecious; epipetric; perennial; rhizomatous.

Rhizome 5-15 cm or more long, (3-)5-10(-16) mm thick,

internodes (2-)5-10(-15) mm long, brown, reddish brown

or greenish brown, sparsely villous. Stipules eventually

deciduous, narrowly to broadly triangular, rarely ovate-

oblong, (3-)5-11 mm long, (3-)4-9 mm wide, greenish,

herbaceous, keeled or weakly keeled, with sparse minute

subsessile brown glandular hairs on abaxial surface,

midrib sparsely villous or glabrous, margin entire, eciliate,

apex aristate, arista 2-3 mm long, hair-like.

Leaves 2-9,

alternate, simple, asymmetric,

unlobed or rarely very

shallowly angular lobed,

suborbicular, base cordate,

margin denticulate and ciliolate, apex obtuse, rounded,

acute or shortly acuminate, (4-)8-14(-25) cm long (basal

lobes included), (3.5-) 7-13(-22) cm wide, adaxially dark

green, brown or dark brown (abaxially reddish or red),

with white maculation (pale green abaxially) along major

veins, sometimes only with some small white dots along

major veins, or pale green along veins, texture thickly

papery, surface rugose, adaxially pilose-setose or villous-

setose (trichomes 1-1.5 mm long, whitish hyaline or

reddish, with a swollen base), abaxially shortly villous,

particularly on veins;

venation basally (6-)7(-8)-palmate,

midrib distinct, veins pinnate along midrib, with 1-3

major lateral veins on each side, other primary veins

branching dichotomously or nearly so, tertiary veins

loosely reticulate and percurrent, forming a divergence

angle of 70-90¢X to major veins, minor veins reticulate, all

veins on abaxial surface prominently raised; petiole terete,

(3-)5-13 cm long, 2-4(-6) mm thick, brownish or brownish

red, villous. Inflorescences axillary, 1-5, dichasial cymes

arising directly from rhizome, 2-4(-6) times-branched,

flowers 5-26(-40); staminate flowers 3-20(-30), carpellate

flowers 2-6(-10); peduncle well developed, taller than

leaves, terete, erect initially, then ascending or pendent,

8-21 cm long, 1.5-2.5 mm thick, brownish, reddish brown

or greenish, sparsely villous or not and with sparse minute

subsessile glandular hairs. Bracts caducous, recurved,

broadly elliptic, oblong or obovate, margin serrulate

and ciliate (with minute glandular tip),

apex obtuse or

nearly rounded, 1.5-11 mm long,

1-7 mm wide, greenish

or brownish, membranaceous, abaxially with minute

subsessile glandular hairs. Staminate flowers: pedicel 6-25

mm long, erect or ascending, with few minute subsessile

glandular hairs; tepals 4, outer 2 suborbicular, oblate or

obovate, base rounded, margin eciliate or occasionally

ciliate below middle, apex rounded, 6-16

mm long,

7-15

mm wide,

pink or pinkish white, with very sparse

minute subsessile brown glandular hairs abaxially; inner

2 tepals oblanceolate, obovate or narrowly so, base

cuneate, margin entire, eciliate, apex obtuse, 6-14 long,

2.5-4 mm wide, white or pinkish, glabrous;

androecium

zygomorphic, tilted, stamens 15-35, golf-club-shaped;

filaments subequal, 1-2 mm long, base slightly fused;

anthers upwardly facing, locules 2, slightly compressed,

oblong-obovoid or narrowly so, apex of connective

emarginate,

1-1.7 mm long, 0.7-0.8 mm wide, yellow.

Carpellate flowers: pedicel 8-17 mm long, horizontal

to pendent, with few minute subsessile glandular hairs,

bracteole absent or rarely with 1 tiny bracteole; tepals 3,

persistent and slightly thickened in fruit; outer 2 tepals

suborbicular or oblate, margin entire, eciliate, 7-11 mm

long, 9-13 mm wide, abaxially pinkish, with sparse minute

subsessile brown glandular hairs; inner tepal narrowly

obovate or obovate, base cuneate, apex obtuse, 5-10 mm

long,

2.5-4 mm wide, pinkish or white, glabrous; ovary

broadly trigonous-ovoid or obovoid, ventral surface

distinctly convex,

6-9 mm long, 4-5 mm thick (wings

excluded), sometimes shortly beaked at apex, greenish

white or reddish white, crisped villous (trichomes 0.5-2

mm long, with a small glandular tip)

and covered with

sparse minute subsessile brown glandular hairs; wings

3, unequal; lateral wings narrower, curved along length

of fruit body,

1-1.5 mm tall; abaxial wing obliquely

triangular or ligulate, rarely obovate-oblong, 3-8 mm tall,

4-6 mm wide; locule 1, placentation intruded parietal

(axile at base);

placentae 3, each 2-branched; styles 3,

slightly tilted, fused at base, apically C-shaped;

stigmatic

band slightly spiraled. Capsule nodding, stipe 8-27 mm

long, greenish or reddish when fresh, 7-11 mm long, 5-8

mm thick (wings excluded), apex with persistent styles

and tepals; lateral wings 1.5-4 mm tall; abaxial wing

obliquely triangular or ligulate, rarely obovate-oblong,

(7-)10-15 mm tall, 5-9(-11) mm wide. Seeds numerous,

brown, ellipsoid or broadly so, 0.45-0.52 mm long,

0.25-0.32 mm thick, chalazal end rounded, micropylar

end obtuse or slightly constricted,

outer periclinal walls

of mature seeds concave; collar cells elongated, straight,

nearly rectangular, (8-)9-12 cells in a ring, occupying

1/4-2/3 of seed length. Somatic chromosome number, 2n =

30 (Figure 3: C).

Additional specimens examined. CHINA. Guangxi

Zhuangzu Autonomous Region: Du¡¦an Xian, Xiaao

Xiang, Jiaba Cun, on shady and moist limestone hill

slope, common, elev. ca. 550 m, 18 Apr 2002, D. Fang &

D. H. Qin 29611 (GXMI); Du¡¦an Xian, Xiaao Xiang,

Guanglong Cun, in a moist cave, elev. ca. 280 m, 11 Jun

2003, D. H. Qin & H. Z. Lin 29616 (GXMI); Longwan

Xiang, on moist limestone hill slope, elev. ca. 200 m,

27 Mar 2003, D. H. Qin & H. Z. Lin 29614 (GXMI);

same locality, 28 Aug 2003, D. Fang & D. H. Qin 29617

(GXMI); Du¡¦an Xian, at entrance of a limestone cave,

24¢X19¡¦12¡¦¡¦N, 107¢X55¡¦01¡¦¡¦E, elev. ca. 440 m scrubby

vegetation, at foot of mountain, on rock face, semishaded,

occasional, C.-I Peng, W. C. Leong, S. M. Ku & Y. Liu

19808 (HAST);

Du¡¦an Xian, Xiaao Xiang, Jiaba Cun, 24¢X

FANG et al. ¡X Three new taxa in

Begonia

sect.

Coelocentrum

in China

107

Figure 7. Begonia retinervia D. Fang, D. H. Qin & C.-I Peng. A, Habit. B, Leaf, cross section. C, Leaf margin, adaxial surface; C',

abaxial surface. D, Stipule, abaxial surface; D', adaxial surface. E, Bract. F, Staminate flower. G, Stamen, lateral view; G', abaxial

view; G", adaxial view. H, Carpellate flower, adaxial view; H', H", lateral views; H"', abaxial view. I, Style and stigmatic band, abaxial

view; I', adaxial view. J, Capsule. K, Dehisced dry capsule. L: a-h, Serial cross sections of ovary. M, Seeds. [A-C, F-H', J from Peng et

al. 19823 (HAST); D from Peng et al. 19808 (HAST); E, H"-I', L, M from Shui 9013; K from Qin & Lin 29614 (GXMI).]

108

Botanical Studies, Vol. 47, 2006

Figure 8. Begonia retinervia D. Fang, D. H. Qin & C.-I Peng. A, Habit and habitat. B, Cultivated plant at anthesis. C, Leaf. D, Leaf,

cross section. E, Leaf margin, adaxial surface. F. Stipule. G, Staminate flower. H, Carpellate flower, lateral view. I, Carpellate flower,

adaxial view. J, Cross section of ovary. K, Capsule. L, Seed SEM microphotograph. [A, F, K from Peng et al. 19808 (HAST); B, D-I

from Peng et al. 19823 (HAST); C, J, L, from Shui 9013 (HAST).]

FANG et al. ¡X Three new taxa in

Begonia

sect.

Coelocentrum

in China

109

14¡¦42¡¦¡¦N, 107¢X55¡¦43¡¦¡¦E, 600 m elev., on limestone rocky

slope, exposed, rare, 31 May 2004, specimens pressed

from plants cultivated in experimental greenhouse at

Academia Sinica, C.-I Peng, W. C. Leong, S. M. Ku &

Y. Liu 19820 (HAST); same locality and date, N-facing

limestone hill, in semi shady depression, specimens

pressed from plants cultivated in experimental greenhouse

at Academia Sinica, C.-I Peng, W. C. Leong, S. M. Ku & Y.

Liu 19823 (HAST); exact locality unknown, 28 May 2002,

Y. M. Shui 9013 (HAST).

Ecology. Rocky limestone slopes and moist

limestone caves, at 200-550 m altitude.

Distribution. North-central Guangxi, China; rare.

(Figure 4).

Phenology. Flowering from August to October;

fruiting from November to March of the next year.

Notes. Begonia retinervia resembles B. ningmingensis,

differing mainly in having leaves thickly papery; ovary

and capsule broadly trigonous-ovoid or obovoid and

crisped villous, with a much-protruded abaxial wing and

markedly convex adaxial side (Table 1). The distribution

ranges of the two species are separated by ca. 200 km.

Acknowledgements. We thank David E. Boufford (A/

GH) and Nicholas J. Turland (MO) for improving the

manuscript; Hidetoshi Nagamasu (KYO) for the Latin

diagnoses; Yan Liu (IBK), Yu-Min Shui (KUN), Wai-Chao

Leong and Huan-Yu Chen (HAST) for accompanying and

assisting us in the field; Hong-Zhe Li (KUN) for providing

a handsome image (Figure 2: A) and the curators of

GXMI, HAST, IBK, IBSC, K, KUN and PE for facilitating

our examination of herbarium collections.

This study was

supported in part by grants from the National Science

Council and Research Center for Biodiversity, Academia

Sinica, Taiwan, to Ching-I Peng (HAST).

Literature Cited

Irmscher, E. 1939. Die Begoniaceen Chinas und ihre Bedeutung

fur die Frage der Formbildung in polymorphen Sippen.

Mitt. Inst. Allg. Bot. Hamburg 10: 431-557.

Ku, T.C. 1999. Begoniaceae. In T. C. Ku (ed.), Flora Reipublicae

Popularis Sinicae, vol. 52(1). Science Press, Beijing, pp.

126-269, Addenda, pp. 401-402.

Ku, S.M., C.-I Peng, and Y. Liu. 2004. Notes on Begonia (sect.

Coelocentrum, Begoniaceae) from Guangxi, China, with

the report of two new species. Bot. Bull. Acad. Sin. 45:

353-367.

Liu, Y., S.M. Ku, and C.-I Peng. 2005. Begonia picturata (sect.

Coelocentrum, Begoniaceae), a new species from limestone

areas in Guangxi, China. Bot. Bull. Acad. Sin. 46: 367-376.

Nguyen, N.Q. 2004. A new species of a peltate-leaved Begonia

(Begoniaceae) from Vietnam. Novon 14: 105-107.

Peng, C.-I, Y.M. Shui, Y. Liu, and S.M. Ku. 2005a. Begonia

fangii (sect. Coelocentrum, Begoniaceae), a new species

from limestone areas in Guangxi, China. Bot. Bull. Acad.

Sin. 46: 83-89.

Peng, C.-I, S.M. Ku, and W.C. Leong. 2005b. Begonia liuyanii

(s ect. Coelocentrum, Begoniaceae), a new species from

limestone areas in Guangxi, China. Bot. Bull. Acad. Sin.

46: 245-254.

S hui, Y.M., C.-I P eng, and C.Y. Wu. 2002. S ynops is of the

Chinense species of Begonia (B egon iac ea e), wi th a

reappraisal of sectional delimitation. Bot. Bull. Acad. Sin.

43: 313-327.

S hu i , Y. M. a n d W.H . Ch e n. 2 0 05 . Ne w d a t a o f s e c t .

Coelocentrum (Begonia) in Begoniaceae. Acta Bot. Yunnan.

27(4): 355-374.

110

Botanical Studies, Vol. 47, 2006

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