ArticlePDF Available

Petrocodon hunanensis (Gesneriaceae), a new species identified by both morphological and molecular evidence from limestone area in Hunan, China

Authors:
  • Guangxi Institute of Botany

Abstract and Figures

Petrocodon hunanensis, a new species of Gesneriaceae from limestone area in Hunan Province, China, is described and illustrated. 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.
Content may be subject to copyright.
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
References
Chen, W.H., Möller, M., Shui, Y.M., Wang, H., Yang, J.B. & Li, G.Y. (2014) Three new species of Petrocodon (Gesneriaceae), endemic
to the limestone areas of Southwest China, and preliminary insights into the diversification patterns of the genus. Systematic Botany
39(1): 316–330.
http://dx.doi.org/10.1600/036364414X678189
Chun, W.Y. (1946) Gesneriacearum novae Sinicarum. Sunyatsenia 6: 271–304.
Fang, D. & Qin, D.H. Wentsaiboea D. Fang & D. H. Qin (2004) A new genus of the
Gesneriaceae from Guangxi, China. Journal of Systematics and Evolution 42 (6): 533–536.
Farris, J.S., Källersjö, M., Kluge, A.G. & Bult, C. (1994) Testing significance of incongruence. Cladistics 10(3): 315–319.
http://dx.doi.org/10.1111/j.1096-0031.1994.tb00181.x
Felsenstein, J. (1985) Confidence limits on phylogenies: an approach using the bootstrap. Evolution 783–791.
http://dx.doi.org/10.2307/2408678
Hance, H.F. (1883) New Chinese Cyrtandreae. Journal of Botany 21: 165–170.
Hemsley, W.B. (1890) Gesneriaceae. In: Forbes, F.B. & Hemsley, W.B. (Eds.) An enumeration of all the plants known from China proper,
Formosa, Hainan, Corea, the Luchu Archipelago, and the Island of Hongkong, together with their distribution and synonymy, part
VIII. Journal of the Linnean Society, Botany 26: 224–234.
http://dx.doi.org/10.1111/j.1095-8339.1890.tb00106.x
Hong, X., Zhou, S.B. & Wen, F. (2014) Petrocodon villosus (Gesneriaceae), a new species from Guangxi, China. Blumea 59: 33–36.
http://dx.doi.org/10.3767/000651914X682378
Jiang, Y.S., Zhang, Y., Wang, Y. & Wen, F. (2011) Petrocodon multiflorus sp. nov. (Gesneriaceae) from Guangxi, China. Nordic Journal
of Botany 29: 57–60.
http://dx.doi.org/10.1111/j.1756-1051.2010.00874.x
Larkin, M.A., Blackshields, G., Brown, N.P., Chenna, R., McGettigan, P. A., McWilliam, H., Valentin, F., Wallace, I.M., Wilm, A., Lopez,
R., Thompson, J.D., Gibson, T.J. & Higgins, D.G. (2007) Clustal W and Clustal X version 2.0. Bioinformatics 23(21): 2947–2948.
http://dx.doi.org/10.1093/bioinformatics/btm404
Li, H.W. (1982) Two new genera and one little known genus of Gesneriaceae from Yunnan. Acta Botanica Yunnanica 4: 241–247.
Li, J.M. & Wang, Y.Z. (2007) Phylogenetic reconstruction among species of Chiritopsis and Chirita sect. Gibbosaccus (Gesneriaceae)
based on nrDNA ITS and cpDNA trnL-F sequences. Systematic Botany 32: 888–898.
http://dx.doi.org/10.1600/036364407783390764
Liu, Y., Xu, W.B. & Huang, Y.S. (2011). Primulina guangxiensis sp. nov. (Gesneriaceae) from a karst cave in Guangxi, China. Nordic
Journal of Botany 29(6): 682–686.
http://dx.doi.org/10.1111/j.1756-1051.2011.01089.x
Liu, Y., Xu, W.B. & Pan, B. (2010) Wentsaiboea tiandengensis sp. nov. and W. luochengensis sp. nov. (Gesneriaceae) from Karst caves in
Guangxi, southern China. Nordic Journal of Botany 28(6): 739–745.
http://dx.doi.org/10.1111/j.1756-1051.2010.00893.x
Möller, M., Forrest, A., Wei, Y.G. & Weber, A. (2011) A molecular phylogenetic assessment of the advanced Asiatic and Malesian
didymocarpoid Gesneriaceae with focus on non-monophyletic and monotypic genera. Plant Systematics and Evolution 292: 223–
248.
http://dx.doi.org/ 10.1007/s00606-010-0413-z
Möller, M., Pfosser, M., Jang, C.G., Mayer, V., Clark, A., Hollingsworth, M.L., Barfuss, M.H.J., Wang, Y.Z., Kiehn, M. & Weber, A. (2009)
A preliminary phylogeny of the ‘Didymocarpoid Gesneriaceae’ based on three molecular data sets: incongruence with available
tribal classifications. American Journal of Botany 96: 989–1010.
http://dx.doi.org/10.3732/ajb.0800291
Nylander, J.A.A. (2004) MrModeltest v2. Program distributed by the author. Evolutionary Biology Centre, Uppsala University, Sweden.
Available from: http://www.downloadatoz.com/developer-johan-nylander.html/ (12 February 2011).
Rogers, S.O. & Bendich, A.J. (1988) Extraction of DNA from plant tissues. In: Gelvin, S.B. & Schilperoort, R.A. (Eds.) Plant Molecular
Biology Manual. Dordrecht: Kluwer Academic Publishers A6.
Ronquist, F. & Huelsenbeck, J.P. (2003) MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 19(12): 1572–
1574.
http://dx.doi.org/10.1093/bioinformatics/btg180
Swofford, D.L. (2002) PAUP*: phylogenetic analysis using parsimony (*and other methods). version 4. Sinauer, Sunderland, Massachusetts.
Taberlet, P., Gielly, L., Pautou, G. & Bouvet, J. (1991) Universal primers for amplification of three non-coding regions of chloroplast
YU ET AL.
72 Phytotaxa 195 (1) © 2015 Magnolia Press
DNA. Plant molecular biology 17(5): 1105–1109.
http://dx.doi.org/10.1007/bf00037152
Wallich, N. (1819) Notice of the progress of botanical science in Bengal. Edinburgh Philosophical Journal 1: 376–381.
Wang, W.T. (1983) Three new genera of Gesneriaceae from China. Bulletin of Botanical Research 1: 15–24.
Wang, W.T. (1984) Notulae de Gesneriaceis Sinensibus VI. Acta Botanica Yunnanica 6: 11–26.
Wang, W.T. & Pan, K.Y. (1982) Notulae de Gesneriaceis Sinensibus (III). Bulletin of Botanical Research 2(2): 121–152.
Wang, Y.Z., Liang, R.H., Wang, B.H., Li, J.M., Qiu, Z.J., Li, Z.Y. & Weber, A. (2010) Origin and phylogenetic relationships of the Old
World Gesneriaceae with actinomorphic flowers inferred from ITS and trnL-trnF sequences. Taxon 59(4): 1044–1052.
Wang, Y.Z., Mao, R.B., Liu, Y., Li, J.M., Dong, Y., Li, Z.Y. & Smith, J.F. (2011) Phylogenetic reconstruction of Chirita and allies
(Gesneriaceae) with taxonomic treatments. Journal of Systematics and Evolution 49: 50–64.
http://dx.doi.org/ 10.1111/j.1759-6831.2010.00113.x
Weber, A., Middleton, D.J., Forrest, A., Kiew, R., Lim, C.L., Rafidah, A.R., Sontag, S., Triboun, P., Wei, Y.G., Yao, T.L. & Möller, M.
(2011b) Molecular systematics and remodelling of Chirita and associated genera (Gesneriaceae). Taxon 60(3): 767–790.
Weber, A., Wei, Y.G., Puglisi, C., Wen, F., Mayer, V. & Möller, M. (2011a) A new definition of the genus Petrocodon (Gesneriaceae).
Phytotaxa 23: 49–67.
Wei, Y.G. (2004) Paralagarosolen Y.G. Wei, a new genus of the Gesneriaceae from Guangxi, China. Journal of Systematics and Evolution
42: 528–532.
Wei, Y.G. (2006) A new species of Lagarosolen W.T. Wang (Gesneriaceae) from Guangxi province, China. Kew bulletin 61: 273–275.
Wei, Y.G., Wen, F., Möller, M., Monro, A., Zhang, Q., Gao, Q., Mou, H.F., Zhong, S.H. & Cui, C. (2010) Gesneriaceae of South China.
Guangxi Science and Technology Publishing House, Nanning, 777 pp.
Wen, F., Liang, G.Y. & Wei, Y.G. (2012) Petrocodon lancifolius (Gesneriaceae), a new species endemic to a central subtropical zone of
Guizhou Province, China. Phytotaxa 49: 45–49.
Wendel, J.F., Schnabel, A. & Seelanan, T. (1995) Bidirectional interlocus concerted evolution following allopolyploid speciation in cotton
(Gossypium). Proceedings of the National Academy of Sciences 92(1): 280–284.
http://dx.doi.org/10.1073/pnas.92.1.280
Xu, W.B., Meng, T., Zhang, Q., Wu, W.H., Liu, Y. & Chung, K.F. (2014) Petrocodon (Gesneriaceae) in the limestone karsts of Guangxi,
China: three new species and a new combination based on morphological and molecular evidence. Systematic Botany 39(3): 965–
974.
http://dx.doi.org/10.1600/036364414X681437
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).
... Wang (2011: 60) has actinomorphic corolla and four or five stamens, while other species have zygomorphic corolla, two stamens and two or three staminodes. Moreover, the color and shape of the corolla now include more variation other than the white, urceolate to campanulate corollas as originally ascribed (Yu et al. 2015). ...
... However, all the plants were rosulate and stemless, different from the caulescent P. hunanensis. We concluded that these plants represent an undescribed species in Petrocodon based on morphological and molecular data and a search through relevant literature (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, Yu et al. 2015. Molecular evidence indicates that this new species, hereafter named P. longitubus Cong R.Li & Yang Luo, is sister to P. hunanensis. ...
Article
Petrocodon longitubus, a new species of Gesneriaceae from Guizhou province, China, is described and illustrated. It is similar to P. hunanensis in having four stamens, but readily differs in being rosulate and stemless and having a narrowly infundibuliform floral tube with the adaxial lip bending upward. Molecular evidence indicates that it is sister to P. hunanensis, but the two species differ markedly in morphology.
... During an expedition in September 2016 investigating the karst flora of Guizhou, we collected a special Petrocodon species in flower from a single locality in Tongzi County, Guizhou, China. The species has four fertile stamens, clearly differentiating it from all described species of Petrocodon except P. hunanensis (Wang et al. 1990, 1998, Wei et al. 2010a, 2010b, Jiang et al. 2011, Liu et al. 2011, Weber et al. 2011, Wen et al. 2012, Hong et al. 2014, Chen et al. 2014, Xu et al. 2014, Li and Wang 2015, Yu et al. 2015, Guo et al. 2016, Cen et al. 2017, Lu et al. 2017. To further investigate the status of this special species and the phylogenetic relevance of the four fertile stamens, we undertook a careful comparison and employed molecular markers to reconstruct the phylogeny of the genus. ...
... ※ indicates the new species, Petrocodon tongziensis. sequence obtained from the author ofYu et al. 2015. ...
Article
Full-text available
Petrocodon tongziensis R.B.Zhang & F.Wen, a new species from northern Guizhou province, China, is described and illustrated based on molecular and morphological evidence. The new species was found growing in crevices and on tufa of moist surfaces of limestone hills in Tongzi County. A maximum parsimony (MP) analysis based on the combined ITS and trnL‐F DNA regions showed that the new species falls within a large polytomy within Petrocodon, but is resolved as most closely related to an unidentified species (WF2014) and these two taxa are in turn resolved as sister to Petrocodon hunanensis X.L.Yu & Ming Li. This is congruent with the fact that both P. hunanensis and P. tongziensis have four fertile stamens, a character state likely to be an ancestral state distinguishing this clade from the rest of Petrocodon. Petrocodon tongziensis differs from P. hunanensis by lacking a terrestrial stem, as well as by the number of bracts, presence of bracteoles, shape of the lobes of the upper lip, and reduced number and length of staminodes.
... Based on the latest molecular phylogenetic analyses, Petrocodon has been expanded to now include a total of about 30 species (Wang et al. 2011, Weber et al. 2011, MÖller et al. 2016, and include all species previously referred to the genera Calcareoboea C.Y. Wu ex H.W. Li, Paralagarosolen Y.G. Wei, Dolicholoma D. Fang & W.T. Wang, Tengia Chun, and Lagarosolen W.T. Wang, plus three species from Didymocarpus Wall., one species from Wentsaiboea D. Fang & D.H. Qin and one species from Primulina Hance (Weber et al. 2011, Wen et al. 2012, Chen et al. 2014, Hong et al. 2014, Xu et al. 2014, Li and Wang 2015, Yu et al. 2015, Guo et al. 2016, IPNI 2016. ...
... Like many new taxa of Petrocodon that have been discovered and described in recent years, P. urceolatus was found in a limestone area and is narrowly endemic. The distribution of Petrocodon species is still restricted to mountainous karst habitats, but has been expanded from south China (Guangdong, Guizhou, Hunan and Guangxi), to southwest China (Yunnan and Congqing), east China (Fujian, Jiangxi) and central China (Hubei) (Chen et al. 2014, Yu et al. 2015, Guo et al. 2016, Qian et al. 2016. This new species is the second new taxa of Petrocodon from Hunan province after P. hunanensis X.L. Yu & Ming Li (Yu et al. 2015). ...
Article
Full-text available
Petrocodon urceolatus F. Wen, H.F. Cen & L.F. Fu, a new species of Gesneriaceae, is described and illustrated from Zhangjiajie city, Hunan Province, China. Morphologically it resembles P. scopulorum (Chun) Y.Z. Wang in the corolla, but differs in several characters including the shape, length and indumentum of the leaf blade, and the length and indumentum of the stamens and staminode. In addition to morphological description, we provide ecological information, geographic distribution and conservation status for this species.
... In contrast, genera like Oreocharis and Petrocodon exhibit a notably high degree of morphological disparity . In particular, both the species diversity and the morphological disparity of the genus Petrocodon have greatly increased since its redefinition in 2011 (Wang et al. 2011;, attributed to the ongoing description of new species in recent years, some of which exhibit a unique floral morphology (Chen et al. 2014;Yu et al. 2015;Guo et al. 2016;Fan et al. 2020;Zhang et al. 2020;Pan et al. 2021). Recently, four species of Allocheilos were reclassified and transferred to the genus Petrocodon , bringing the total number of species in this genus to 54, including one variety. ...
Article
Full-text available
The genus Petrocodon is renowned for its remarkably diverse floral morphology, exhibiting a high level of disparity compared to most genera within Gesneriaceae. In this study, we present a detailed description and illustration of Petrocodon gracilis T. Ding & B. Pan, a new species with unique floral features that is native to Guangxi, China. This species is geographically close to P. jingxiensis H.S. Gao & W.B. Xu and is indistinguishable from P. jingxiensis in vegetative characteristics. However, it can be clearly distinguished from the latter by its shorter pedicels, slender tubular corolla, shorter corolla tube, distinctly unequal corolla lobes with upper ones linear and reflexed and lower ones oblanceolate, and included chiritoid‐like stigma. The presence of linear upper lobes and the chiritoid‐like stigma has not been previously reported in Petrocodon, and the emergence of this new species will further expand the morphospace occupied by this morphologically diverse genus.
... Recircumscriptions have led to quite radical redefinitions and/or expansions of some genera (e.g., Deinostigma, Oreocharis, Petrocodon, and Primulina). In Petrocodon, which until recently was known only to include one penta-and more than 20 diandrous species, now also includes three tetrandrous species (Yu et al., 2015;Li et al., 2019;Zhang et al., 2019). Evidently, much work is still necessary before we have a satisfactory understanding of all genera to then be reflected in a satisfactory classification. ...
... We carefully observed the morphology of those plants that were introduced and cultivated in the nursery and common garden of GCCC over the following years and confirmed that it can be clearly distinguished from Petrocodon integrifolius. Detailed analysis of morphology, types and other authentic specimens of related species of Petrocodon housed in various herbaria (E, GH, HN, IBK, K, KUN, MO, PE, PH, US and VNMN) and relevant literature (Wang et al., 1990(Wang et al., , 1998Li & Wang, 2005;Wei, 2006Wei, , 2007Wei et al., 2010b;Wen et al., 2012;Chen et al., 2014;Jiang et al., 2011;Hong et al., 2014;Xu et al., 2014;Li & Wang, 2015;Middleton et al., 2015b;Yu et al., 2015;Guo et al., 2016;Lu et al., 2017b;Cen et al., 2017, Zhang et al., 2018Li et al., 2019;Su et al., 2019 a & b;Zhang et al., 2019) revealed it to be a new a taxon, which is described and illustrated here. Perennial herbs forming acaulescent rosettes. ...
Article
Full-text available
A new species, Petrocodon ionophyllus F.Wen, S.Li & B.Pan, from the limestone area of Southwestern Guangxi, South China is described here. Based on the population information and threatened situation, the current existence status of this species is assessed as 'Endangered'.
... To test the systematic placement of the new species, the nuclear ribosome internal transcribed spacers (ITS) region and chloroplast trnL-F intron spacer (trnL-F) were chosen to reconstruct the phylogenetic tree of Petrocodon, since these two regions were frequently used in previous taxonomic and phylogenetic works of (e.g. Möller et al. 2009, 2011, Chen et al. 2014, Xu et al. 2014, Yu et al. 2015, Lu et al. 2017a, 2017b, Zhang et al. 2018, 2019. We extracted total genomic DNA of the new species and Petrocodon hunanensis from silica-dried leaves collected from the field using a CTAB method (cf. ...
... Möller & A.Weber in Weber et al. (2011a: 782) and P. pinnata (W.T. Wang (1984a: 25) Y.Z. Wang in Wang et al. (2011: 62) as outgroup based on recent phylogenetic analyses (Möller et al. 2009, Weber et al. 2011a& 2011b, Yu et al., 2015. Sequences obtained from this study and NCBI GenBank are listed in Table 1. ...
Article
Full-text available
Petrocodon asterocalyx F.Wen, Y.G.Wei & R.L.Zhang, a new species from the Danxia landform area in Guangxi, South China, is described and illustrated based on molecular and morphological data. The molecular evidence shows that the new species is recovered in a weakly supported clade. Within this clade, the new one is morphologically similar to P. hancei (Hemsl.) A.Weber & Mich.Möller and P. coriaceifolius (Y.G.Wei) Y.G.Wei & Mich.Möller, and it can be distinguished from the former by calyx lobes 20–40 × 2–3 mm, corolla 2.5–3.0 cm long, filaments sparsely erectly pubescent, anthers sparsely pubescent, staminodes 3, and stigmas 2; from latter by leaf blades rhombic-oblong or rhombic, base shallowly cuneate, margin crenulate to serrate, calyx lobes linear, 20–40 × 2–3 cm, and anthers 3.5–3.8 mm long, sparsely pubescent and elliptical.
Article
Full-text available
A new species, Petrocodon chishuiensis Z.B. Xin, F. Wen & S.B. Zhou, from NW Guizhou, China, is described and illustrated here. It shares a rare characteristic with P. hunanensis X. Luo: all have four fertile stamens, a character state distinguishing this clade from the rest of Petrocodon. P. chishuiensis closely resembles P. hunanensis and P. tongziensis, but differs in vegetative and generative characters. We found only one population with no more than 60 mature individuals at the type locality. This species is provisionally assessed as Critically Endangered (CR B2ab(iii)) using IUCN criteria.
Article
Full-text available
Based on molecular studies, the small Chinese genus Petrocodon (two species and one variety) has been recently enlarged to include the monotypic genera Calcareoboea, Paralagarosolen and Tengia. It is shown here that the (6-7) species of Lagarosolen, the monotypic Dolicholoma, a few species of Didymocarpus, and a number of new species that have recently been published (but not formally described) under Petrocodon and Lagarosolen should be included in this genus. This raises the size of the genus from five to around 20 species. With respect to the floral diversity (corolla form, size, and coloration; with the exception of Tengia, the androecium is always diandrous) and inferred pollination syndromes (different forms of melittophily, ornithophily, psycho- and/or sphingophily), Petrocodon represents one of the most varied genera of Old World Gesneriaceae, comparable to some New World genera.
Article
Full-text available
A new species of Gesneriaceae from southern China, Petrocodon lancifolius, is described and illustrated. It is similar to another stenochoric species, P. scopulorum, but is differentiated from its congeners by several characters, such as the shape of the leaf blades, the indumentum of peduncle, pedicels, the outside of bracts and corolla, the corolla shape, limb and lobes, the ovary, pistil, style, and phenology. A description of P. lancifolius, together with illustrations, a distribution map, habitat description, conservation assessment and a diagnostic comparison with P. scopulorum are presented.
Article
Full-text available
A new species of Gesneriaceae from south-western Guangxi, China, Petrocodon villosus, is described and illustrated. It resembles P. ferrugineus, but is differentiated by several characters, such as the size of bracts, calyx and corolla, the indumentum of leaf blades, calyx, filaments and the outside of the corolla; the corolla tube shape, limb and lobes, the disc and stigma. A description of P. villosus, together with illustrations, habitat description and a diagnostic comparison are presented.
Article
The phylogenetic placement of the Old World Gesneriaceae genera Ramonda, Conandron, Bournea, Thamnocharis, and Tengia, all characterized by actinomorphic flowers, has been the subject of much debate. Actinomorphy in Gesneriaceae is rare, with most species exhibiting zygomorphic flowers. The actinomorphic genera have historically been considered “primitive” and lumped in the tribe Ramondeae separate from the remaining Old World Gesneriaceae. In this study, we used nuclear (ITS) and plastid (trnL‐F) DNA for molecular phylogenetic analysis of these five genera along with representative species across the Cyrtandroideae. Our results show that the actinomorphic genera are scattered over several otherwise zygomorphic clades within Cyrtandroideae, and along with previous data, indicate that Ramondeae is an unnatural group. Floral actinomorphy has evolved convergently in different alliances of Old World Gesneriaceae. Ramonda is sister to Haberlea, Bournea is apparently paraphyletic, Conandron seems rather isolated, and Tengia is close to Petrocodon and sister to a group of Chirita sect. Gibbosaccus together with Calcareoboea. We hypothesize that the evolution from zygomorphy to actinomorphy with novel combinations of characters is possibly due to shifts in pollination strategies, such as a switch from nectar‐ to pollen‐rewards.
Article
A new species of Lagarosolen, L. coriaceifolium Y. G. Wei, from Guangxi province, China is described and illustrated. Its relationship with Lagarosolen integrifolius D. Fang & L. Zeng is discussed.
Article
Wentsaiboea renifolia D. Fang & D. H. Qin, a new genus and species of Gesneriaceae from Guangxi, China, is described and illustrated. This genus is similar to Dolicholoma D. Fang & W. T. Wang in the shape of stigma, but differs by having leaves reniform, palmately nerved and base cordate, corolla obliquely campanulate, corolla lobes rounded, and both stamens and staminodes adnate to corolla tube near base. It is also similar to Chiritopsis W. T. Wang in the habit, but differs by having leaves palmately nerved, corolla tube campanulate and abaxially swollen, and stigma hippocrepiform.
Article
Sequences from the internal transcribed spacer region of nuclear DNA (ITS) and the trnL-F regions of chloroplast DNA for fifteen species of Chirita and seven species of Chiritopsis were used to assess phylogenetic relationships between Chiritopsis and Chirita section Gibbosaccus. Parsimony and Bayesian inference (BI) analyses were conducted using separate nuclear and chloroplast data sets, as well as a combined data set. Phylogenetic trees resulting from separate analyses proved highly congruent and the combined analysis of the two data sets produced a well-supported topology of the species and sections examined. Section Gibbosaccus proved paraphyletic and Chiritopsis polyphyletic in all analyses. Our results provide evidence that the species of Chiritopsis are embedded in section Gibbosaccus and were derived at least twice from within section Gibbosaccus. Finally, we reconsider the morphological evolution and adaptation between and within the two genera. The present analyses indicate that nomenclatural changes will be needed to reflect more accurately relationships in the Gibbosaccus-Chiritopsis complex. Only about 10% of the species in Chirita have been sampled and further data are required before any taxonomic changes can be suggested.
Article
Paralagarosolen fangianum Y. G. Wei, a new genus and species of Gesneriaceae from Guangxi, China, is described and illustrated. Paralagarosolen Y. G. Wei is closely related to Lagarosolen W. T. Wang in having corolla tube cylindric, not swollen, and stigmas 2, but differs by having leaves sometimes peltate at base, cyme with only one flower, corolla lobes rounded-obtuse at apex, and capsule broadly ovoid-ellipsoid.