Content uploaded by Shuai Peng
Author content
All content in this area was uploaded by Shuai Peng on May 17, 2021
Content may be subject to copyright.
Phytotaxa 500 (3): 217–224
https://www.mapress.com/j/pt/
Copyright © 2021 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Accepted by Prabhukumar K.M.: 28 Apr. 2021; published: 13 May 2021
https://doi.org/10.11646/phytotaxa.500.3.5
217
Impatiens bullatisepala (Balsaminaceae), a new species from Guizhou, China
SHUAI PENG1,2,3,4,5, YI-YAN CONG1,6*, JING TIAN2,3,7, CAI-FEI ZHANG2,3,8, GUANG-WAN HU2,3,9* & QING-
FENG WANG2,3,10
1 College of Life Sciences, Hunan Normal University, Changsha 410081, China
2 CAS Key Laboratory of Plant Germplasm Enhancement and Specialty Agriculture, Wuhan Botanical Garden, Chinese Academy of
Sciences, Wuhan 430074, China
3 Sino-Africa Joint Research Center, Chinese Academy of Sciences, Wuhan 430074, China
4 University of Chinese Academy of Sciences, Beijing, 100049, China
5
�
pengshuai183@163.com; https://orcid.org/0000-0002-8064-920X
6
�
908158033@qq.com; https://orcid.org/0000-0001-8900-1111
7
�
tianjing@wbgcas.cn; https://orcid.org/0000-0002-6716-7513
8
�
zhangcf@wbgcas.cn; https://orcid.org/0000-0002-2818-5751
9
�
guangwanhu@wbgcas.cn; https://orcid.org/0000-0001-7728-7976
10
�
qfwang@wbgcas.cn; https://orcid.org/0000-0001-9143-8849
*Corresponding Authors
Abstract
Impatiens bullatisepala (Balsaminaceae), a new species supported by morphological and phylogenetic evidence from Fanjing
Mountain, Guizhou province in China, is described here. It is morphologically similar to I. davidii but can be distinguished
by its dorsally ridged lateral sepals with sunk reticulate veins and bullate projections on abaxial surface, 2–2.5 cm deep
saccate lower sepal with ca. 0.8 cm long narrowly triangular tip at the mouth, and broadly ovate dorsal petal. Phylogenetic
analyses of a combined dataset of nuclear ITS and plastid atpB-rbcL intergenic spacer DNA sequences furtherly confirmed
its novelty.
Keywords: Impatiens davidii, new taxon, taxonomy
Introduction
The genus Impatiens Linnaeus (1753: 937) (Balsaminaceae) consist of more than 1000 species is one of the most
species-rich genera in angiosperms and mainly distributed in tropical and subtropical Africa and Eurasia (Grey-Wilson
1980, Yuan et al. 2004, Yu et al. 2016). Yu et al. (2016) divided this genus into two subgenera: I. subgen. Impatiens
S.X. Yu & Wei Wang (2016: 191) and I. subgen. Clavicarpa S. X. Yu ex S. X. Yu & Wei Wang (2016:191) based on
molecular and morphological evidence. I. subgen. Impatiens is further divided into seven sections, of them I. sect.
Impatiens is characterized by its 1- or 2-flowered racemose inflorescences, linear or cylindrical capsule, and ellipsoid
seed. In recent years, a lot of new species have been described (Kuang et al. 2014, Cai et al. 2015, Tan et al. 2015,
Ding et al. 2016, 2017, Gan et al. 2016, Guo et al. 2016, Wang et al. 2016) on account of extensive field investigation
and careful study of the botanist. This has brought the number of Impatiens species in China to more than 280 (Chen
et al. 2007, Du et al. 2020).
In October 2006, during a field trip on Fanjing Mountain in Guizhou province, one of the authors (G.-W. Hu)
came across an interesting Impatiens along the side of a creek under evergreen broad-leaved forests. Its lateral sepals
with sunk reticulate veins and bullate projections on the abaxial surface made it quite different from other Impatiens
species. The primary studies could not give this species a correct scientific name. To correctly identify it, another
field survey was carried out at the same place and its neighboring areas in September 2018. Then, we collected some
specimens for further research. After consulting relevant literature (Chen et al. 2001, 2007, Xiong et al. 2009, Yu et al.
2012) and new taxa described in recent years, we found that Xiong & Yang (2009) identified it as I. dicentra Franchet
ex Hook. f. (1908: 268) in the book named Illustrated Handbook of Common Herbaceous Seed Plants in Fanjing
PENG ET AL.
218 • Phytotaxa 500 (3) © 2021 Magnolia Press
Mountain National Nature Reserve. However, we found it was morphologically different to I. dicentra, notably in the
lateral sepals. I. dicentra owns the lateral sepals with coarsely dentate margin, flat and smooth abaxial surface, while
this particular species owns the lateral sepals with entire margins and sunk reticulate veins and bullate projections
on abaxial surface. After comparing the specimens in PE, KUN, HIB, and HNNU and studying in morphology and
molecular phylogenetics, we concluded that it is a new species in the Impatiens sect. Impatiens and describe it here as
I. bullatisepala sp. nov.
TABLE 1. GenBank accession numbers for samples in the phylogenetic analysis.
Species ITS atpB-rbcL
Impatiens piufanensis KP776094
Impatiens imbecilla AY348796 DQ147851
Impatiens lateristachys KP776078 KP776030
Impatiens faberi AY348778 DQ147841
Impatiens forrestii AY348784 DQ147847
Impatiens oxyanthera AY348814 DQ147865
Impatiens microstachys KP776085
Impatiens neglecta KP776087 KP776038
Impatiens pterosepala KP776097 KP776046
Impatiens macrovexilla KP776082 KP776034
Impatiens leptocaulon KP776080 KP776032
Impatiens capensis AY348759 DQ147823
Impatiens noli-tangere KP776088 KP776039
Impatiens tortisepala KP776106 KP776054
Impatiens fissicornis AY348782 DQ147844
Impatiens lecomtei AY348802 DQ147855
Impatiens platychlaena AY348818 DQ147867
Impatiens soulieana AY348833 DQ147880
Impatiens barbata AY348750 DQ147818
Impatiens corchorifolia AY348767 DQ147831
Impatiens chiulungensis KP776066 KP776016
Impatiens delavayi AY348773 DQ147836
Impatiens nubigena KP776089 KP776040
Impatiens poculifer AY348820 DQ147870
Impatiens chekiangensis KP776064 KP776014
Impatiens platysepala KP776095 KP776044
Impatiens textorii AY348841
Impatiens tienmushanica KP776105 KP776053
Hydrocera triflora AY348853 DQ147895
A NEW SPECIES OF IMPATIENS BULLATISEPALA Phytotaxa 500 (3) © 2021 Magnolia Press • 219
FIGURE 1. Impatiens bullatisepala A. Plant in nature habit; B. Flower bud; C. Flower lateral view; D. Flower front view; E. Different
parts of a flower; F. Fruit. Scale bars: B–F = 1 cm. Photographed by Shuai Peng.
PENG ET AL.
220 • Phytotaxa 500 (3) © 2021 Magnolia Press
Materials and methods
Morphologic description and comparison
The morphological description of the new species was based on the examination of fresh material in the field and
herbarium specimens. Comparisons with other species were referenced to herbarium specimens kept in KIB, HNNU,
HIB, and PE, photographs, and literature (Grey-Wilson 1980, Chen et al. 2001, Chen et al. 2007, Yu et al. 2012).
Phylogenetic analyses
To speculate the phylogenetic status of the new species, one nuclear gene region: ITS (Yuan et al. 2004), and one
plastid sequence: atpB-rbcL (Janssens et al. 2006) were used in the phylogenetic analysis. We used three individuals
of the new species (voucher specimens: PS-0052-a, PS-0052-b & PS-0052-c) and six individuals from two populations
of Impatiens davidii Franchet (1886: 65) (voucher specimens: HGW-001138, HGW-001140, HGW-001142, PS-0051-a,
PS-0051-b & PS-0051-c) for phylogenetic analyses. Voucher specimens were deposited in the herbarium of Wuhan
Botanical Garden, CAS (HIB). Genomic DNA of new species was extracted from silica gel dried leaves with Mag-
MK Plant Genomic DNA Extraction Kits (Sangon Biotech, Shanghai). PCR products were sequenced by TSINGKE
Biological Technology.
The sequences of 28 taxa in Impatiens sect. Impatiens and the outgroup taxon were acquired from GenBank
(Assession numbers provided in Table 1). Phylogenetic reconstruction was performed using Bayesian inference (BI).
MEGA v.7.0 (Kumar et al. 2016) was used to align nuclear and plastid sequences, then manually integrate. MrModeltest
v.2.3 (Nylander 2004) was used to determine the best models of sequence evolution for a combined dataset using the
Akaike Information Criterion (ITS + atpB-rbcL: GTR + I + G). Bayesian majority rule consensus trees of datasets
were constructed by MrBayes v.3.2.6 (Ronquist et al. 2012). The dataset was run for 10,000,000 generations, starting
from different random trees and sampled every 100 generations. The initial 25% of sampled trees were discarded as
burn-in.
Taxonomy
Impatiens bullatisepala G.W. Hu, Y.Y. Cong & Q.F. Wang sp. nov. (Figures 1, 2: A–D)
Diagnosis:—The new species is morphologically similar to Impatiens davidii, but can be distinguished with the latter by its lateral sepals
suborbicular, 1–1.5 cm in diameter, midrib ridged dorsally, lateral veins reticulate and sunk on the abaxial surface with bullate
projections among veins; dorsal petal broadly ovate, 1–1.5 × 1.5–2 cm, dorsally with a green narrow arcuate cristate crest.
Type:—CHINA. Guizhou province, Jiangkou County, Fanjing Mountain, under evergreen broad-leaved forest, 27°53′44.21″N,
108°43′16.4″E, elevation 1027 m, 15 September 2018, Shuai Peng PS-0052 (holotype: HIB!; Isotype: HIB!, HNNU!)
Herbs annual, to 100 cm tall. Stem erect or procumbent in lower part, succulent, branched, glabrous, lower nodes
swollen with many fibrous roots. Leaves alternate, petiolate; petiole 0.5–2 cm long; lamina elliptic to elliptic-lanceolate,
6–10 × 3–4.5 cm, margin coarsely crenate, teeth mucronulate, base obtuse to cuneate, with 3–4 pairs of glands, apex
caudate-acuminate; glabrous; lateral veins 5–7 pairs. Inflorescences in upper leaf axils, 1-flowered; peduncle 1.8–2.8
cm long. Pedicel with 2 bracts above middle; bracts alternate, glabrous, upper one ovate, ca. 6 × ca. 4 mm, lower one
subulate, ca. 2–3 mm long. Flower yellow, 4–5.5 cm long. Lateral sepals 2, sub-orbicular, 1–1.5 cm in diameter, apex
mucronulate, margin entire, yellowish-green, glabrous, midrib ridged dorsally, lateral veins reticulate and sunk on
abaxial surface with bullate projections among veins, ; lower sepal deeply saccate, 2–2.5 cm deep, yellow with 6–7
orange red striations, glabrous; apex gradually narrowed into an abruptly incurved spur, spur 2-lobed, 1–1.3 cm long;
mouth 1.5–1.9 cm wide, oblique with ca. 0.8 cm long narrowly triangular tip. Dorsal petal broadly ovate, 1–1.5 × 1.5–2
cm, apex emarginate and mucronulate, dorsally with a green narrow arcuate cristate crest at midvein; lateral united
petals 2-lobed, 2.4–3.2 cm long, basal lobe ovate to elliptic, 5–7 × 3–5 mm, small, upper part with purple-red patches,
apex abruptly narrowed into a purple-red filiform appendage, appendage up to 1 cm long; distal lobe dolabriform,
1.8–2.3 × 0.9–1.4 cm, yellow with red spots at base, inner margin curled with small auricle, apex with a short yellow
filiform appendage. Stamens up to 4 mm long, filaments linear, slightly swollen above; anthers ovoid-orbicular, apex
obtuse. Ovary erect, fusiform. Capsule linear, 2.8–3.5 cm long. Seed unknown.
A NEW SPECIES OF IMPATIENS BULLATISEPALA Phytotaxa 500 (3) © 2021 Magnolia Press • 221
FIGURE 2. Impatiens bullatisepala A–D: A. Flower bud; B. Flower lateral view; C. Flower front view; D. Different parts of a flower.
Impatiens davidii E–K: E & I. Flower bud; F & J. Flower lateral view; G. Flower front view; H & K. Different parts of a flower. Scale
bars: A–K = 1 cm. A–D & I–K: photographed by Shuai Peng, E–H: photographed by Guang-Wan Hu (A–D from S. Peng PS-0052 in HIB,
E–H from G.-W. Hu & S. Peng HGW-001138 in HIB, I–K from S. Peng PS-0051 in HIB).
Distribution and ecology:—Impatiens bullatisepala is currently known only from the type locality in Fanjing
Mountain, Guizhou Province, China. It grows under the moist evergreen broad-leaved forests along the side of a
creek.
Etymology:—The specific epithet “bullatisepala” refers to the lateral sepals with bullate projections netted in
sunk reticulate veins on abaxial surface. The Chinese name is “泡萼凤仙花”.
Phenology:—The new species was observed flowering from September to October, fruiting from October to
November.
Phylogenetic position:—Bayesian phylogenetic analyses of the combined ITS and atpB-rbcL datasets proved its
position within the Impatiens sect. Impatiens and closely related to I. davidii in the phylogenetic tree (Figure 3).
Voucher specimens of Impatiens davidii:—China. Hunan: Yongshun County, Xiaoxi National Natural Reserve,
28°48′N, 110°11′E, elevation 650 m, 15 August 2018, S. Peng PS-0051 (HIB); Jiangxi: Jiujiang City, Lushan Mountain,
Guling Township, 29°34′N, 115°58′E, elevation 1050 m, 11 October 2018, G.W. Hu & S. Peng HGW-001138, G.W. Hu
& S. Peng HGW-001140, G.W. Hu & S. Peng HGW-001142 (HIB).
PENG ET AL.
222 • Phytotaxa 500 (3) © 2021 Magnolia Press
FIGURE 3. Bayesian phylogenetic tree base on a combined dataset of nuclear ITS and plastid atpB-rbcL DNA sequences, in which
Impatiens bullatisepala is showing in red, and Bayesian posterior probabilities are indicated for each branch. (I. davidi1 from HGW-
001138, I. davidii2 from HGW-001140, I. davidii3 from HGW-001142, I. davidii4–6 from PS-0051, all specimens deposited in HIB)
Discussion
In Impatiens sect. Impatiens, a group of closest relatives similar to I. davidii including I. fissicornis Maximowicz
(1889: 87), I. platychlaena Hook. f. (1908: 270), I. lecomtei Hook. f. (1908: 271) and I. dicentra. They are showing a
high similarity in having 1- or 2-flowered raceme inflorescence with yellow flowers, saccate lower sepal with several
striations, short incurved spur, and the lateral united petal with a long filamentous appendage, but mainly differ in
the shape of lateral sepals, dorsal petal, and spur (Chen 2001). Based on morphological characteristics, we found that
there are continuous variations within this group. Therefore, this poses challenges to the classification of species in
this group. Moreover, the evidence from the combinations of morphology and molecules in this particular group may
be more convincing for taxonomy to species. More detailed comparisons among the new species and its close relatives
in I. sect. Impatiens present in Table 2.
Acknowledgments
We acknowledge the National Wild Plant Germplasm Resource Center for all kinds of support. This research was
supported by the International Partnership Program of Chinese Academy of Sciences (No. 151853KYSB20190027)
and National Natural Science Foundation of China (No. 31970211).
A NEW SPECIES OF IMPATIENS BULLATISEPALA Phytotaxa 500 (3) © 2021 Magnolia Press • 223
TABLE 2. Comparison of floral morphological characters of Impatiens bullatisepala to I. davidii and other closely related
Impatiens spp.
Characters I. bullatisepala I. davidii I. fissicornis I. platychlaena I. lecomtei
Leaves
elliptic to elliptic-
lanceolate, 6–10 ×
3–4.5 cm
elliptic, 5–10 × 3–4
cm
ovate-oblong or
ovate-lanceolate,
4–7 × 2–3 cm
ovate-oblong, ovate,
or ovate-lanceolate,
6–15 × 3–4.5 cm
ovate or ovate-
lanceolate, or
oblong-ovate, 10–15
× 2.5-3 cm
Flower yellow yellowish yellow or orange-
yellow
bicolored: purple and
yellow pink
Lateral sepal
Sub-orbicular,
dorsal ridged
and vesicular
protuberances,
margin entire
broadly ovate,
dorsal 9-veined
subovate-orbicular,
dorsal
orbicular, dorsal
many veined, mid-
vein not thickened
orbicular dorsal
mid-vein carinate,
7-veined
Lower sepal deeply saccate saccate saccate deeply saccate broadly funnelform
Spur incurved hooked hooked incurved incurved
Dorsal petal
broadly ovate,
dorsally with a
narrow arcuate
cristate crest at
mid-vein
sub-orbicular,
abaxial mid-vein
cristate
sub-orbicular,
abaxial mid-vein
broadly thickened,
cristate
orbicular,
mucronulate, abaxial
mid-vein thickened,
broadly cristate
orbicular, abaxial
mid-vein thickened,
broadly cristate,
crista entire
References
Cai, X.Z., Hu, G.W. & Cong, Y.Y. (2015) Impatiens xanthinoides (Balsaminaceae), a new species from Yunnan, China. Phytotaxa 227
(3): 261–267.
https://doi.org/10.11646/phytotaxa.227.3.5
Chen, Y.L. (2001) Balsaminaceae. In: Chen, Y.L. (Ed.) Flora Reipublicae Popularis Sinica, vol. 47 (part 2). Science Press, Beijing, pp.
1–243.
Chen, Y.L., Akiyama, S. & Ohba, H. (2007) Balsaminaceae. In: Wu, Z.Y. & Raven, P.H. (Eds.) Flora of China, vol. 12. Science Press,
Beijing & Missouri Botanical Garden Press, St. Louis, pp. 43–113.
Ding, B., Gadagkar, S.R., Wang, J.C., Zhang, M., Guo, H. & Yu, S.X. (2016) Impatiens wawuensis (Balsaminaceae): a new species from
Sichuan, China. Phytotaxa 273 (4): 293–298.
https://doi.org/10.11646/phytotaxa.273.4.5
Ding, B., Wang, J.C., Deng, H.P. & Wang, C.Y. (2017) Impatiens baishaensis (Balsaminaceae): a new species from Sichuan, China.
Phytotaxa 319 (2): 191–196.
https://doi.org/10.11646/phytotaxa.319.2.8
Du, C., Liao, S., Boufford, D.E. & Ma, J.S. (2020) Twenty years of Chinese vascular plant novelties, 2000 through 2019. Plant diversity
42 (5): 393–398.
https://doi.org/10.1016/j.pld.2020.08.004
Franchet, M.A. (1886) Plantae Davidianae ex sinarum imperio: Par MA Franchet 1. G. Masson, Paris, 485 pp.
Gan, Q.L. & Li, X.W. (2016) Impatiens baokangensis (Balsaminaceae), a new species from Hubei, China. Annales Botanici Fennici 53:
145–148.
https://doi.org/10.5735/085.053.0401
Guo, H., Zhang, C.F., Zhang, L.J. & Yu, S.X. (2016) Impatiens guiqingensis (Balsaminaceae), a new species from Gansu, China. Phytotaxa
247 (3): 229–233.
https://doi.org/10.11646/phytotaxa.247.3.6
Grey-Wilson, C. (1980) Impatiens of Africa. Balkema, Rotterdam, 235 pp.
PENG ET AL.
224 • Phytotaxa 500 (3) © 2021 Magnolia Press
Hooker, J.D. (1908) Les Espèces Du Genre “Impatiens” Dans L’herbier Du Muséum De Paris. Nouvelles archives du Muséum d’histoire
naturelle sér. 4 (10): 233–272.
Janssens, S.B., Geuten, K., Yuan, Y.M., Küpfer, P. & Smets, E.F. (2006) Phylogenetics of Impatiens and Hydrocera using plastid atpB-
rbcL spacer sequences. Systematic Botany 31: 171–180.
https://doi.org/10.1600/036364406775971796
Kuang, R.P., Duan, L.D., Gu, J.Z., Cai, X.Z., Cong, Y.Y. & Liu, K.M. (2014) Impatiens liboensis sp. nov. (Balsaminaceae) from Guizhou,
China. Nordic Journal of Botany 32 (4): 463–467.
https://doi.org/10.1111/j.1756-1051.2013.00194.x
Kumar, S., Stecher, G. & Tamura, K. (2016) Mega 7: molecular evolutionary genetics analysis version 7.0 for bigger datasets. Molecular
Biology & Evolution 33 (7): 1870.
https://doi.org/10.1093/molbev/msw054
Linnaeus, C. (1753) Species Plantarum, vol. 2. Salvius, Stockholm, 1200 pp.
https://doi.org/10.5962/bhl.title.669
Maximowicz, C.J. (1889) Plantae Chinenses Potaninianae nec non Piasezkianae. Trudy Imperatorskago S.-Peterburgskago Botaničeskago
Sada 11: 1-112.
Nylander, J.A.A. (2004) MrModeltest (Version 2.3): Evolutionary Biology Centre, Uppsala University. [https://github.com/nylander/
MrModeltes2]
Ronquist, F., Teslenko, M., vander Mark, P., Ayres, D.L., Darling, A., Hohna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck, J.P.
(2012) MrBayse 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61
(3): 539–542.
https://doi.org/10.1093/sysbio/sys029
Tan, Y.H., Liu, Y.N., Jiang, H., Zhu, X.X., Zhang, W. & Yu, S.X. (2015) Impatiens pandurata (Balsaminaceae), a new species from
Yunnan, China. Botanical studies 56 (1): 29.
https://doi.org/10.1186/s40529-015-0108-4
Wang, Q., Gadagkar, S.R., Deng, H.P., Yang, Z.M. & Yu, F.Q. (2016) Impatiens shennongensis (Balsaminaceae): a new species from
Hubei, China. Phytotaxa 244 (1): 96–100.
http://dx.doi.org/10.11646/phytotaxa.244.1.8
Xiong, Y.X. & Yang, C.D. (2009) Illustrated Handbook of Common Herbaceous Seed Plants in Fanjing Mountain National Nature
Reserve. Guizhou Science and Technology Publishing House Co., Ltd, Guiyang, pp. 91–95.
Yu, S.X. (2012) Balsaminaceae of China. Peking University Press, Beijing, 215 pp.
Yu, S.X., Janssens, S.B., Zhu, X.Y., Lidén, M., Gao, T.G. & Wang, W. (2016) Phylogeny of Impatiens (Balsaminaceae): integrating
molecular and morphological evidence into a new classification. Cladistics 32 (2016): 179–197.
https://doi.org/10.1111/cla.12119
Yuan, Y.M., Song, Y., Geuten, K., Rahelivololona, E., Wohlhauser, S., Fischer, E., Smets, E. & Küpfer, P. (2004) Phylogeny and
biogeography of Balsaminaceae inferred from ITS sequence data. Taxon 53: 391–404.
https://doi.org/10.2307/4135617
