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
pg_0002
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.]
pg_0003
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).]
pg_0004
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
pg_0005
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.
pg_0006
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.]
pg_0007
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.]
pg_0008
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
pg_0009
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
pg_0010
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
pg_0011
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).]
pg_0012
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).]
pg_0013
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.
pg_0014
110
Botanical Studies, Vol. 47, 2006
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