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Phytotaxa 195 (1): 065–072
www.mapress.com/phytotaxa/
Copyright © 2015 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Accepted by Zhi-Qiang Zhang: 10 Dec. 2014; published: 22 Jan. 2015
http://dx.doi.org/10.11646/phytotaxa.195.1.4
65
Petrocodon hunanensis (Gesneriaceae), a new species identified by both
morphological and molecular evidence from limestone area in Hunan, China
XUN-LIN YU1*, MING LI1, JIAN-JUN ZHOU1 & PENG-WEI LI2
1The Dendrological Teaching and Research Team, School of Forestry, Central South University of Forestry & Technology, CN-410004,
No. 498, Shaoshan South road, Changsha city, Hunan province, China. E-mail: csfuyuxl@163.com *(author for corresponding), limin-
gjx@126.com, zhoujianjun00188@126.com
2State key Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences, Xiangshan Road, Nanx-
incun 20, Beijing 100093, China. E-mail: pdlc@163.com
Abstract
Petrocodon hunanensis, a new species of Gesneriaceae from limestone area in Hunan Province, China, is described and il-
lustrated. The new species is morphologically similar to Petrocodon coriaceifolius, but readily differs from the latter one in
the growth form with terrestrial stems and distinct internodes, both surfaces of leaf densely with white pubescence, petiole
densely with reddish-purple or white pubescence, zygomorphic corolla white or pale purple and 2–3 cm long, pedicels 0.3–2
(2.8) cm long, 4 stamens and 1 staminodes, ovary and capsule stipitate. Molecular evidences indicate that it is systematically
similar to P. hispidus, but the morphologies of two relatives are obviously different.
Key words: Calcareoboea, Didymocarpus, Dolicholoma, Lagarosolen, limestone flora, molecular evidence, new species,
Paralagarosolen, Petrocodon, Tengia, Wentsaiboea
Introduction
Based on the recent molecular studies (Wang et al. 2010; Möller et al. 2011; Wang et al. 2011; Weber et al. 2011a,
2011b; Xu et al. 2014), several genera have been integrated into Petrocodon Hance (1883: 167), including the
monotypic genera Calcareoboea C.Y. Wu ex H.W. Li (1982: 241), Paralagarosolen Y.G. Wei (2004: 528), Tengia
Chun (1946: 279), Dolicholoma D. Fang & W.T. Wang in Wang (1983: 18), all the species of Lagarosolen W.T. Wang
(1984: 11), one species of Wentsaiboea D. Fang & D.H. Qin (2004: 533) (Wentsaiboea tiandengensis Yan Liu & B.
Pan 2010: 739), three species of Didymocarpus Wall. (1819: 378) (Didymocarpus niveolanosus D. Fang & W.T. Wang
in Wang & Pan 1982: 133, Didymocarpus mollifolius W.T. Wang 1984: 21, Didymocarpus hancei Hemsl. 1890: 229)
and one species of Primulina Hance (1883: 169) (Primulina guangxiensis Yan Liu & W.B. Xu in Liu et al. 2011: 682).
Consequently, the number of species in the genus Petrocodon increased to 26, including nine new species published
recently, mainly distributed in southern China and northern Vietnam (Wei et al. 2010; Jiang et al. 2011; Weber et al.
2011a; Wen et al. 2012; Chen et al. 2014; Hong et al. 2014; Xu et al. 2014). The re-circumscription of Petrocodon
increased the morphological variability of this genus (Weber et al. 2011a). For instance, Petrocodon scopulorus (Chun)
Y.Z. Wang has an actinomorphic corolla with 4 or 5 stamens, while the other species have zygomorphic corollas and 2
stamens and 2 or 3 staminodes. Moreover, the corolla color and shape now include more types compared to the white
and urceolate to campanulate forms of original Petrocodon.
During field investigations of the limestone areas of Hunan Province in 2013 and 2014, we collected plants with a
corolla shape and leaf blade shape most similar to Petrocodon coriaceifolius (Y.G. Wei 2006: 273) Y.G. Wei & Mich.
Möller in Weber et al. (2011a: 59). However, these plants have 4 stamens and 1 staminode which readily differentiates
it from P. coriaceifolius (with 2 stamens and 2 staminodes). Eventually, we confirm it as a new species of the newly re-
circumscribed genus, Petrocodon based on the morphological and molecular data and some relevant literatures (Wang
1984; Wei 2006; Wei et al. 2010; Jiang et al. 2011; Weber et al. 2011a; Wen et al. 2012; Chen et al. 2014; Hong et al.
2014; Xu et al. 2014;). Molecular evidences indicate that it is systematically similar to P. hispidus (W.T. Wang 1984:
12) A. Weber & Mich. Möller in Weber et al. (2011a: 60), but the morphologies of these two relatives are obviously
YU ET AL.
66 • Phytotaxa 195 (1) © 2015 Magnolia Press
different. This new species is a unique species in Petrocodon for its 4 fertile stamens in a zygomorphic corolla, the
terrestrial stem with distinct internodes and distinctly stipitate ovary and capsule. The new species, Petrocodon huna-
nensis X. L. Yu & Ming Li, is described and illustrated below.
Material and Methods
Plant matierials
Thirteen species were sampled and combined with the new species. Most of the ITS and trnL-F DNA sequences were
acquired from GenBank (Appendix 1). Leaf materials of three relative species (Petrocondon hunanensis sp. nov., P.
mollifolius, Didymocarpus yunnanensis) for newly acquired sequences were collected in the field and dried by silica
gel (Appendix 2). Voucher specimens were deposited in PE and CSFI.
The ingroup comprises eight species of Petrocondon (P. coriaceifolius, P. dealbatus, P. hancei, P. hispidus, P.
mollifolius, P. niveolanosus, P. scopulorum, P. hunanensis sp. nov.). The outgroup comprises six species, including
two species of Chirita (C. anachoreta, C. pumila), two species of Didymocarpus (D. cortusifolius, D. yunnanensis),
and two species of Primulina (P. heterotricha, P. tabacum) (Li & Wang 2007; Möller et al. 2009; Wang et al. 2011;
Weber et al. 2011a, 2011b).
DNA extraction and PCR
The total genomic DNA which used as the template in the polymerase chain reaction (PCR) was extracted from
silica gel dried leaves and used the modified CTAB method of Rogers & Bendich (1988). The ITS and trnL-F were
amplified using the ITS primers ITS1 and ITS4 (Wendel et al. 1995) and the trnL-F primers c and f (Taberlet et al.
1991), respectively.
Sequence alignment and Phylogenetic analysis
The sequences were aligned using Clustal W (Larkin et al. 2007) and adjusted manually in the software Geneious
version 7.1.4.
The incongruence length difference test (Farris et al. 1994) was implemented in PAUP*4.0B10 (Swofford 2003)
to assess potential congruence between ITS and trnL-F and the p value (0.979) showed the two regions matched each
other greatly. Thus, we concatenated them sequentially to carry out the followed analysis.
Parsimony analysis for the combined matrices was implemented in PAUP*4.0B10 using maximum parsimony
methods. All characters were unordered type and had equal weight. Gaps were treated as missing data. Starting trees
obtained via stepwise addition. Heuristic searches were carried out with 1000 replicates of random addition, one tree
was held at each step during stepwise addition, branch-swapping strategy was tree-bisection-reconnection (TBR). In
order to evaluate the robustness of different clades, we conducted a bootstrap analysis (Felsenstein 1985) with 1000
replicates of bootstrapping using a heuristic search with 1000 replicates of random sequence addition and TBR branch
swapping.
To select a suitable model of sequence evolution, the combined matrices were computed in Mrmodeltest version
2.3 (Nylander 2004) and the result suggested the GTR+I+G model best fitted the combined data. Bayesian inference
analyses were carried out in MrBayes version 3.2.2 (Ronquist & Huelsenbeck 2003). 1 000 000 generations were run
in two independent analyses each with four Markov chain Monte carlo (MCMC) chains. One tree was sampled every
500 generations (=2000 trees) and the first 400 trees discarded as burn-in. Posterior probabilities (PP) obtained from
the analysis were used to indicate the credibility of various branches.
Results and Discussion
Parsimony analyses resulted in eight trees of equal length (L=649; CI=0.795; RI=0.655). The MP tree comprises four
clades and each clade represents a genus (Fig. 1).
PETROCODON HUNANENSIS (GESNERIACEAE) Phytotaxa 195 (1) © 2015 Magnolia Press • 67
FIGURE 1. One of eight most parsimonious trees generated from analysis of combined ITS and trnL-F data for all sampled taxa.
Bootstrap (BS) values are on the left side of the slash and Bayesian posterior probabilities (PP) on the right. The asterisk indicates no
support. The bold indicates the new species, Petrocodon hunanensis X. L. Yu & Ming Li.
The new species undoubtedly locates in strongly supported clade IV (BS=95%, PP=99%), which exclusively
consists of taxa from Petrocodon (Fig. 1). Phylogenetically, the new species is sister with Petrocodon hispidus but with
relatively low support (BS=50%, PP=99%). Moreover, the morphologies of these two relative species are obviously
different.
YU ET AL.
68 • Phytotaxa 195 (1) © 2015 Magnolia Press
Taxonomy treatment
Petrocodon hunanensis X. L. Yu & Ming Li, sp. nov. (See Figs. 2 and 3).
Diagnose:—The new species differs from Petrocodon coriaceifolius (Y.G. Wei) Y.G. Wei & Mich. Möller by having subterete terrestrial
stem with distinct internodes, both surface of leaf densely with white pubescence, petiole densely with reddish-purple or white
pubescence, corolla white to pale purple and 2–3 cm long, tube infundibuliform, pedicels 0.3–2 (2.8) cm long, densely pubescent,
stamens 4 and staminodes 1, ovary and capsule stipitate.
Type:—CHINA. Hunan Province: Dong’an Xian, Zixishi Zhen, Huzhu Village [26°17′N, 111°15′E], elevation ca. 230 m, on the rock
surface or at the entrance of karst caves. 28 August 2014, J.J. Zhou 14082801 (holotype CSFI!, isotypes CSFI!)
Perennial herbs, terrestrial stem subterete, with distinct internodes, 5–20 cm long, ca. 5 mm in diameter. Leaves
crowded at the top of the stem; leaf blade thickly papery, ovate, 2–3.2 × 1–2.5 cm, apex slightly acuminate to obtuse,
base cuneate to broadly cuneate, leaf blade adaxial surface green and abaxial surface reddish-purple (or rarely pale
green), both surfaces densely white pubescent, margin crenate, petiole 0.5–2.5 cm long, densely with reddish-purple or
white pubescence; lateral veins 4 pairs, adaxially impressed, abaxially prominent. Cymes 1–4, 1–4-flowered; peduncles
FIGURE 2. Petrocodon hunanensis X. L. Yu & Ming Li, sp. nov. (Drawn by Jing Tian): (A) plant with flowers, (B) calyx, (C) opened
corolla showing stamens and staminodes, (D) pistil, (E) capsule.
PETROCODON HUNANENSIS (GESNERIACEAE) Phytotaxa 195 (1) © 2015 Magnolia Press • 69
FIGURE 3. Petrocodon hunanensis X. L. Yu & Ming Li, sp. nov.: (a) habitat, (b) and (c) mature plant in the wild, (d) lateral view of
flower, (e) opened corolla showing stamens and staminodes, (f) pistil, (g) calyx and disc, (h) seed, (i) young fruit.
YU ET AL.
70 • Phytotaxa 195 (1) © 2015 Magnolia Press
2–7 cm long, densely reddish-purple pubescent; bracts 2, opposite, ovate-lanceolate, 1–3 × 1 mm, apex obtuse, margin
entire, outside densely villous, inside with sparse pubescence. Pedicels 0.3–2 (2.8) cm long, densely pubescent. Calyx
5-parted to the base, lobes triangular-lanceolate, 3–5 × 1 mm, densely with short puberulence except the base of inside
glabrous, margin entire. Corolla white to pale purple, 2–3 cm long, outside densely puberulent, inside pubescent and
with short glandular hairs, tube infundibuliform, 0.8–1.2 cm in diameter at the mouth, 1.5–2 cm long, inside with
several pale purple stripes; limb 2-lipped, adaxial lip ca. 4 mm long, 2-lobed, lobes ovate, abaxial lip ca. 1.2 cm long,
3-lobed, central one longer, long ovate, 6–7 mm long, ca. 4 mm in diam., lateral ones diagonal ovate or triangular
ovate, ca. 6mm long and 5 mm in diam.. Two pale yellow patches at the base of the central abaxial lip lobe, densely
with short glandular hairs. Stamens 4 (coherent in pairs of two), two of them longer, adnate to ca. 1.1 cm above the
base of the corolla tube, filaments 0.7–1 cm long, the other two adnate to ca. 0.9 cm above the base of the corolla tube,
filaments 0.5–0.7 cm long, and all filaments densely with short glandular hairs and pubescence; anthers elliptic, ca. 1.5
mm long, with short glandular hairs and pubescence, pollen chambers 4; staminodes 1, adnate to ca. 0.6 cm above the
base of corolla tube, ca. 1 mm long, glabrous. Pistil ca. 1.6 cm long, ovary distinctly stipitate and ca. 1 cm long, style
0.6–0.8 cm long, all densely with glandular hairs and pubescence; stigma 2, ovate, ca. 1 mm long, densely pubescent.
Disc annular, yellow, ca. 1 mm in height, margin entire. Capsule linear and distinctly stipitate, 4-valved, 2.5–4 cm ×
1–2 mm, with densely short puberulence. Seeds black, elongated or spindle.
Distribution and habitat:—The new species is only known from its type locality, namely Dong’an Xian, Zixishi
Zhen, Huzhu Village, Hunan Province, China. It grows on moist shady rock surfaces or at the entrance of a limestone
hill, at an elevation of ca. 230 m.
Phenology:—Flowering from July to September, and fruiting from August to October.
Etymology:—The specific epithet is derived from the type locality of the new species, i.e. Hunan Province,
China.
Similar species:—Petrocodon hunanensis is morphologically similar to Petrocodon coriaceifolius in having
similar leaf blade shape and corolla shape, but it differs from P. coriaceifolius by having subterete terrestrial stem
with distinct internodes, both surfaces of leaf densely with white pubescence, petiole densely with reddish-purple or
white pubescence, corolla white to pale purple, tube infundibuliform and 0.8–1.2 cm in diam. at the mouth, pedicels
0.3–2 (2.8) cm long, densely pubescent, stamens 4 and staminodes 1, ovary ca. 1 cm long, capsule densely with short
puberulence and ovary and capsule distinctly stipitate. The detailed contrasting characters between two relative species
are shown in Table 1.
TABLE 1. Morphological comparison between Petrocodon hunanensis and P. coriaceifolius
Character P. hunanensis P. coriaceifolius
Terrestrial stem with distinct internodes, 5–20 cm long, 5 mm in diam. no
Leaf blade both surface of leaf densely with white pubescence with appressed white pubescence on each side,
with purple glands on the dorsal surface
Cymes 1–4, 1–4-flowered 1–2, 7–14-flowered
Petiole 0.5–2.5 cm long, densely with reddish-purple or white pubescence 0.5–5.4 cm long, densely with white pubescent
Pedicel 0.3–2 (2.8) cm long 0.3–0.5 cm long
Corolla color white to pale purple pale purple
Corolla tube infundibuliform, 0.8–1.2 cm in diam. at mouth, 1.5–2 cm long slender, 0.5 cm in diam. at mouth, 1.2–1.3 cm
long
Stamens 4, filaments 0.7–1 cm long 2, filaments 0.5–0.6 cm long
Staminodes 1, ca. 1 mm long 3, 0.1–0.2 mm long
Ovary ca. 1 cm long, stipitate ca.0.8 cm long
Capsule stipitate and densely with short puberulence densely pubescent
Ackonwlegments
The authors are grateful to Jing Tian for preparing the line drawing.
PETROCODON HUNANENSIS (GESNERIACEAE) Phytotaxa 195 (1) © 2015 Magnolia Press • 71
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Appendix 1. GenBank accession numbers (Species: trnL-F / ITS).
Petrocodon coriaceifolius (Y.G. Wei) & Mich. Möller: HQ632943/ HQ633040; Petrocodon dealbatus Hance:
FJ501537/ FJ501358; Petrocodon hancei (Hemsl.) A. Weber & Mich. Möller: HQ632944/ HQ633041; Petrocodon
hispidus (W.T. Wang) A. Weber & Mich. Möller: HQ632939/ HQ633036; Petrocodon niveolanosus (D. Fang & W.T.
Wang) A. Weber & Mich. Möller: JF697588/ JF697576; Petrocodon scopulorum (Chun) A. Weber & Mich. Möller:
HQ632947/ HQ633044; Chirita anachoreta Hance: DQ872820/ DQ872837; Chirita pumila D. Don: FJ501491/
FJ501327; Didymocarpus cortusifolius (Hance) Levl.: HQ632898/ HQ632995; Primulina heterotricha (Merr.) Y.Z.
Wang: DQ872816/ DQ872826; Primulina tabacum Hance: AJ492300/ FJ501352.
Appendix 2. Voucher with collection locality and herbarium where deposited in.
Petrocodon hunanensis X. L. Yu & Ming Li: J.J. Zhou 14082801, Hunan, China (CSFI); Petrocodon mollifolius (W.T.
Wang) A. Weber & Mich. Möller: LJM 2012001, Yunnan, China (PE); Didymocarpus yunnanensis (Franch.) W.W.
Smith: LPW 2012028, Sichuan, China (PE).