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Swertia subuniflora (Gentianaceae), a new species from Fujian, China

Phytotaxa 280 (1): 036–044
Copyright © 2016 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
36 Accepted by Hai He: 24 Sept. 2016; published: 14 Oct. 2016
Swertia subuniflora (Gentianaceae), a new species from Fujian, China
1 College of Life Science, Fujian Normal University, Fuzhou 350117, China
2 Key laboratory of Adaption and Evolution of Plateau Biota, Northwest Institute of Plateau Biology, Chinese Academy of Sciences, Xin-
ing 810001, China; E-mail:
3 University of Chinese Academy of Sciences, Beijing 100039, China.
These authors contributed equally to this work
Swertia subuniflora is described from Fujian Province, China. Morphological and molecular evidences indicate its close
relationship with S. franchetiana, S. mussotii and S. punicea, three species from Swertia series Japonica (further grouped in
section Swertopsis, subgenus Ophelia). The new species can be recognized by its solitary flowers and ovate-cordate stem
Keywords: cpDNA, morphology, new taxon, taxonomy
Swertia Linnaeus (1753: 226) is one of the largest genera of Gentianaceae, with a wide distribution in Asia, North
America, Europe and Africa (Ho 1988, Ho & Pringle 1995). The genus contains over 160 species, which have been
grouped into three subgenera and eleven sections (Ho 1988, Rybczyński et al. 2014, Ho & Liu 2015a). There are 75
species in China according to Flora of China (Ho & Pringle 1995).
The new species described here first came to our attention in October 2010 during our Mountain Jinrao expedition,
which was initially misidentified as Lomatogonium macranthum (Diels & Gilg 1903: 17) Fernald (1919: 197) (Lin et
al. 2014). Subsequent study of floral morphology, however, indicated that this plant was actually a member of Swertia.
In September 2014, with additional flowering specimens available, and combined with newly generated molecular
data, its identity was further confirmed as a new species of Swertia. And hereafter it is described as Swertia subuniflora
B.H.Chen & S.L.Chen.
TABLE 1. Morphological comparisons among Swertia subuniflora, S. mussotii, S. punicea, S. franchetiana and S. cordata.
S. subuniflora S. punicea S. mussotii S. franchetiana S. cordata
stem leaves ovate-cordate,
4–12 × 3–8 mm
lanceolate, or narrowly
to 60 × 18 mm
narrowly lanceolate to
8–35 × 3–10 mm
lanceolate to ovate-
15–37 × 2–8 mm,
8–23 × 5–12 mm
inflorescence branched proximally,
each branch 1-flowered
many flowered
paniculate cyme
many flowered
paniculate cyme
many flowered
paniculate cyme
flower merosity 5 usually 5, occasionally 4 4 5 5
corolla diameter equal in size
about 10 mm
unequal in size,
8–12 mm
equal in size
8–13 mm
equal in size
15–25 mm
equal in size
10–15 mm
corolla color white with blue veined
yellow or dark purple dark purple pale blue pale purple with dark
purple veins
oblong, radially
elongated, boat-shaped,
two nectaries per corolla
radially elongated, with
a narrow scale, raised
margin with many long
fimbriae, two nectaries
per corolla lobe
radially elongated, with
a narrow scale, raised
margin with many long
fimbriae, two nectaries
per corolla lobe
radially elongated, with
a narrow scale, raised
margin with many long
fimbriae two nectaries
per corolla lobe
rhomboid to orbicular
nectary, reduced to a
naked gland patch, one
nectary per corolla lobe
SWERTIA SUBUNIFLORA (GENTIANACEAE) Phytotaxa 280 (1) © 2016 Magnolia Press 37
Materials and methods
Leaf material of S. subuniflora for DNA extraction was collected as described by Gao et al. (2012). Voucher specimens
were deposited in HNWP.
The treatment of leaves and DNA extraction were carried out following the method of Fu et al. (2016). The plastid
genes matK and rbcL were amplified and sequenced according to the protocols described by Sun et al. (2013).
Parallel chromatograms derived from bi-directional sequencing were checked for accuracy by visual inspection
with Chromas v. 2.33 ( and integrated into a single sequence. The sequences were
then aligned and trimmed according to those of Swertia species taken from GenBank. The two sequences for S.
subuniflora have been submitted to GenBank (Accession numbers KX019837–KX019842, also see Table 2). They
were then concatenated for phylogenetic analysis.
Phylogenetic relationships of S. subuniflora, with 37 other species of Swertia, representing eight sections, especially
Swertia sect. Swertopsis (Makino 1901: 47) Satake (1947: 25), were resolved by means of maximum parsimony (MP)
as implemented in PAUP v. 4.0b10 (version for 32bit Microsoft Windows, Swofford 2003). Four species of Gentiana
Linnaeus (1753: 227) and three of Lomatogonium A. Braun (1830: 221) in Gentianaceae were utilized as outgroups.
Parameters for MP analysis were set following Li et al. (2016).
TABLE 2. Taxa, references (or vouchers) and GenBank accession numbers of Gentiana and Swertia used in present study.
An asterisk (*) denotes sequences newly obtained for this study. A line (—) denotes sequences not obtained.
Species Reference or Vouchers
Accession (matK) Reference or Vouchers
Accession (rbcL)
G. crassicaulis Xi et al., 2014 KC861277.1 Malik et al., 2015 KJ947517.1
G. straminea Xi et al., 2014 KC861281.1 Chen et al., 2010 GQ436509
G. dahurica Xi et al., 2014 KC861279.1 Unpublished KM226715.1
G. macrophylla Chen et al., 2010 GQ434141.1 Unpublished KM226708.1
L. rotatum Sun et al., 2013 KC935910.1 Sun et al., 2013 KC935898.1
L. rotatum var. tenuifolium Saarela et al., 2013 KC474959.1 Saarela et al., 2013 KC483095.1
L. macranthum Xi et al., 2014 KC861247.1 HNWP, LiuJQ-263 KX645868*
S. parryi von Hagen et al., 2002 AJ408029.1 & AJ408022.1 — —
S. tubulosa von Hagen et al., 2002 AJ408030.1 & AJ408023.1 — —
S. bifolia Xi et al., 2014 KC861232.1 HNWP, 25151 KX019852*
S. marginata Xi et al., 2014 KC861235.1 HNWP, 0463 KX019847*
S. wolfangiana Xi et al., 2014 KC861227.1 HNWP, 26118 KX019843*
S. erythrosticta Xi et al., 2014 KC861266.1 HNWP, 005 KX019849*
S. gyacaensis Xi et al., 2014 KC861248.1 — —
S. wardii Xi et al., 2014 KC861244.1 HNWP, 7622 KX019845*
S. perennis Thiv et al., 1999 AJ010528.1 & AJ011457.1 — —
S. cordata Unpublished KC594672.1 HNWP, PAK8998 KX019844*
S. angustifolia Kshirsagar et al., 2015 LM644053.1 Kshirsagar et al., 2015 LM644059.1
S. angustifolia var. pulchella Kshirsagar et al., 2015 HG917352.1 Kshirsagar et al., 2015 HG964542.1
S. corymbosa Kshirsagar et al., 2015 HG917351.1 Kshirsagar et al., 2015 HG964541.1
S. densifolia Kshirsagar et al., 2015 HG917339.1 Kshirsagar et al., 2015 HG964529.1
S. lawii Kshirsagar et al., 2015 HG917350.1 Kshirsagar et al., 2015 HG964539.1
S. nervosa Xi et al., 2014, KC935915.1 Kshirsagar et al., 2015 LM644055.1
S. abyssinica Struwe et al., 2002 AJ388191.1 & AJ388261.1 — —
S. crassiuscula Hagen et al., 2001 AJ406347.1 & AJ406375.1 — —
S. alba Sun et al., 2013 KC935911.1 Sun et al., 2013 KC935899.1
S. macrosperma CPBOL Group et al., 2011 JF956570.1 CPBOL Group et al., 2011 JF944536.1
...continued on the next page
38 Phytotaxa 280 (1) © 2016 Magnolia Press
TABLE 2. (Continued)
Species Reference or Vouchers
Accession (matK) Reference or Vouchers
Accession (rbcL)
S. franchetiana CPBOL Group et al., 2011 JF956568.1 CPBOL Group et al., 2011 JF944534.1
S. mussotii Xi et al., 2014 KC861255.1 HNWP, 391 KX019846*
S. punicea CPBOL Group et al., 2011 JF956579.1 CPBOL Group et al., 2011 JF944543.1
S. subuniflora 1 HNWP, B.H.Chen 092201 KX019837* HNWP, B.H.Chen 092201 KX019840*
S. subuniflora 2 HNWP, B.H.Chen 092202 KX019838* HNWP, B.H.Chen 092202 KX019841*
S. subuniflora 3 HNWP, B.H.Chen 092203 KX019839* HNWP, B.H.Chen 092203 KX019842*
S. chirayita Kshirsagar et al., 2015 HG917353.1 Kshirsagar et al., 2015 HG964543.1
S. cincta Xi et al., 2014 KC861259.1 Xi et al., 2014 JF944525.1
S. dilatata Xi et al., 2014 KC861265.1 HNWP, 7615 KX019850*
S. pubescens CPBOL Group et al., 2011 JF956575.1 CPBOL Group et al., 2011 JF944541.1
S. racemosa Xi et al., 2014, KC861263.1 — —
S. hispidicalyx Unpublished KP184452.1 Unpublished KP184457.1
S. tenuis CPBOL Group et al., 2011 JF956582.1 CPBOL Group et al., 2011 JF944547.1
S. yunnanensis CPBOL Group et al., 2011 JF956584.1 CPBOL Group et al., 2011 JF944553.1
S. leducii Sun et al., 2013 KC861268.1 Sun et al., 2013 KC935903.1
S. rosularis Xi et al., 2014 KC861251.1 — —
S. patens Xi et al., 2014 KC861233.1 — —
S. bimaculata CPBOL Group et al., 2011 JF956556.1 CPBOL Group et al., 2011 JF944518.1
S. paniculata Kshirsagar et al., 2015 LM644051.1 Kshirsagar et al., 2015 LM644057.1
S. dichotoma Hagen et al., 2002 AJ408035.1 & AJ408028.1 HNWP, 1033 KX019851*
The MP tree (Fig. 3) showed that S. subuniflora has a very close phylogenetic relationship with S. franchetiana H.Smith
(1907: 251), S. mussotii Franchet (1899: 316) and S. punicea Hemsley in Forbes & Hemsley (1890: 140), all belonging
to Swertia subgen. Ophelia C.B.Clarke (1883: 129), sect. Swertopsis, ser. Japonica T.N.He & S.W.Liu (2013: 389).
This indicates that S. subuniflora may also be a member of ser. Japonica. It is worth noting that the present defined
subgenus Ophelia is paraphyletic with respect to other subgenera and to the genus Lomatogonium. This situation was
also recognized in previous studies (He et al. 2013a, b, Yuan et al. 2003, Xi et al. 2014). The relationship between
Swertia and its allies needs to be further clarified (Ho & Liu 2015b, Jiu-Li Wang unpublished data).
Swertia subuniflora B.H.Chen & S.L.Chen, sp. nov. (Figs. 1 & 2)
Type:—CHINA. Fujian: Jianning County, Mount Jinrao, on wet grassland slopes or beside streams, 116°55’38” E, 26°45’9” N, elev. 1798
m, 22 September 2014, B.H. Chen 092201 (holotype, FNU!; isotypes, HNWP!, FNU!).
Description:—Herbs annual, 10–35 cm tall. Roots yellow, stout. Stems erect, subquadrangular, with narrow wings on
angles; basal branches approximately isometric, erect, ca. 1 mm in diameter; upper branches anisometric, ascending,
slender. Basal leaves spatulate, withered at anthesis, long petiolate; stem leaves, ovate-cordate, sessile, 4–12 × 3–8
mm, apex obtuse, veins pinnate, only mid-vein distinct, base auriculate and subamplexicaul; distal leaves gradually
smaller. Flowers 5-merous, 10 mm in diam., solitary and terminal on each flowering stem, ranging from (1–) 2–10
(–23) per plant. Pedicel 7–30 mm, erect, slender. Calyx green, 1/3–1/2 as long as corolla; lobes ovate-lanceolate, 5–8
SWERTIA SUBUNIFLORA (GENTIANACEAE) Phytotaxa 280 (1) © 2016 Magnolia Press 39
FIGURE 1. Swertia subuniflora. A. Plant with flower (bud). B. Flower. C. Petals imbricate in bud. D. Stamens. E. Corolla and stamens.
F. Petal. G. Pistil and calyx. H. Pistil and stigma. I. Longitudinal section of ovary.
40 Phytotaxa 280 (1) © 2016 Magnolia Press
FIGURE 2. Swertia subuniflora. A–C. Plants and habitat. D–E. Complete plant. F. calyx. G. A collora lobe. H. Filament position. I–J.
Stamens. K. Pistil. L. Longitudinal section of ovary. M. Stigma.
SWERTIA SUBUNIFLORA (GENTIANACEAE) Phytotaxa 280 (1) © 2016 Magnolia Press 41
FIGURE 3. Phylogenetic relationships resulting from analysis of matK+rbcL sequences in Swertia by maximum parsimony (MP). I.
Outgroups. II. Swertia subgen. Poephila. III. Lomatogonium. IV. Swertia subgen. Ophelia. V. Swertia subgen. Swertia.
42 Phytotaxa 280 (1) © 2016 Magnolia Press
× 2–4 mm, apex acuminate, abaxially 1-veined. Corolla white with blue-veins, membranous, petals imbricate in bud;
lobes ovate to ovate-lanceolate, 11–13 × 4–5 mm, apex obtuse. Nectaries 2 per corolla lobe, oblong, radially elongated,
boat-shaped, margin fimbriate, 3–4 mm long. Filaments linear, ca. 4.8 mm, not dilated at base; anther violet, oblong,
ca. 2 mm. Ovary sessile, ovate, surface covered with purple spots, ca. 5 mm; style short and salient; stigma lobes
Distribution and habitat:—Swertia subuniflora is so far only known from the type locality in Jianning County,
Fujian Province, China. It grows on wet grassland slopes or along streams at elevations of 1700–1800 m, in association
with plants such as Pinus taiwanensis Hayata (Pinaceae), Eurya saxicola Hung T. Chang (Pentaphylacaceae),
Stranvaesia davidiana var. undulata (Decaisne) Rehder & E. H. Wilson (Rosaceae), Ligularia japonica (Thunberg)
Lessing (Asteraceae), Gentiana davidii Franchet (Gentianaceae), Astilbe macrocarpa Knoll (Saxifragaceae),
Miscanthus sinensis Andersson (Poaceae) and Molinia japonica Hackel (Poaceae).
Phenology:—Flowering and fruiting from September to October.
Etymology:—The specific epithet refers to the flower which is almost solitary on each flowering stem.
Relationships:—Swertia subuniflora is an annual with fibrous roots, the basal leaves are few and deciduous
at anthesis and stem leaves are ovate-cordate (Fig. 1A, Fig. 2D & E). Flowers are 5-merous (Fig. 2B & C). Stamen
filaments are not dilated at the base (Fig. 2J) and are clearly separated from each other (Fig. 2H & I). The two nectaries
of each corolla lobe are straight, opening from base of corolla (Fig. 2G). With these features the new species resembles
S. punicea, S. franchetiana and other species from Swertia ser. Japonica, this relationship is also supported by plastid
sequences in the MP analysis. However, the new species can be easily distinguished by leaf and floral morphology
(Table 1).
From its overall vegetative appearance, especially the ovate-cordate stem leaves, Swertia subuniflora is also
similar to S. cordata (Wallich ex G. Don 1837: 178) C.B.Clarke (1883: 123). However, it can be distinguished by its
unique inflorescence and floral characters (Table 1).
Although the new species had originally been misidentified as Lomatogonium macranthum (Lin et al. 2014), the
latter is distinguished by bicolored flowers and decurrent stigma. Their separation is also confirmed by the MP analysis
(Fig. 3).
Additional specimens examined (paratypes):—CHINA. Fujian: Jianning County, Jinrao Mountain. 22
September 2014, B.H.Chen 092202 (HNWP!); B.H.Chen 092203 (FNU!); 3 October 2010, Y.L. Qiu, C.N. Fang &
B.H. Chen 2010100303 (FNU!).
We thank Ms. Yanlian Qiu, Mr. Yunlong Gao and Mr. Weiwen Song for their kind help during the field work, and
Ms. Taili Cai for preparing the line drawing. We are very grateful to Dr. R. J. Gornall for polishing English of the
manuscript. This work was funded by the National Natural Science Foundation of China (Grants No. 31270270 &
31400322), Sub-project VI of National Program on Key Basic Research Project (Grant No. 2015FY110200), CAS
“Light of West China” Program, Youth Innovation Promotion Association CAS, and the international scientific and
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... Moreover, several new species have been described from China since the account of the genus for the Flora of China was published (e.g. Chen et al. 2008;Liu 2010, 2015;Chen et al. 2016). ...
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The genetic structure and phylogeographical history of the alpine shrubs Sibiraea angustata (Rosaceae) and Sibiraea laevigata from the Qinghai–Tibetan Plateau (QTP) were investigated to identify alpine plant responses to changes in the QTP and glaciations. Fifty-five populations were analyzed using four chloroplast DNA (cpDNA) regions and (nuclear ribosomal internal transcribed spacer) nrITS sequence data. In all, 21 cpDNA haplotypes and 13 nrITS sequence types were detected. Analyses of the genetic diversity and phylogenetic relationships detected two rarely reported glacial refugia. One was the Yushu–Nangqian area, and the other consisted of the area from the Songpan Plateau to the southeastern margin of the QTP. Sibiraea species populations experienced divergent evolution and founder effects when they recolonized the QTP platform and adjacent high-altitude regions following glaciations. The divergence times of the main lineages and haplotypes were in the range of 1.60–2.58 Ma. The population size of Sibiraea species in the QTP decreased approximately 23-fold during the last 0.12 Ma, indicating that Sibiraea species were significantly affected by environmental changes in the QTP. Therefore, the rapid uplift of the QTP and subsequent glaciations likely played an important role in driving genetic divergence and population size changes of Sibiraea species in the QTP.
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We evaluated the phylogeography and historical demography of the cyprinid fish Gymnodiptychus dybowskii (subfamily Schizothoracinae) across three northern Qinghai-Tibetan Plateau (QTP) river systems in the Tien Shan range: the Kaidu River, Ili River and Junggar Basin. Results from both mtDNA (16S rRNA and Cyt b) and nuDNA (RAG-2) resolved three reciprocally monophyletic clades, one in each of the three river basins. Estimated divergence times (highest posterior density (HPD) 2.4-3.7 Mya) are consistent with the hypothesis that these three clades are products of vicariance resulting from the intensive uplift of QTP and Tien Shan, and resulting expansion of the Taklimakan and Gurbantunggut deserts. Several lines of evidence indicate dynamic demographic histories for the three clades, with late Quaternary population bottlenecks and expansions in the Kaidu and Ili rivers and, possibly, a Holocene decline in the Junggar Basin. For conservation purposes, the three clades should be treated as species or minimally, as evolutionarily significant units (ESUs). They have experienced decades of anthropogenic disturbance and preservation of the three species/ESUs will require more sustainable management of the aquatic resources.
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Swertia is ethno-medicinally an important genus belonging to family Gentianaceae. Swertia chirayita is used as imperative medicinal plant in Indian system of medicine. However, this species has been frequently adulterated due to its high demand and scarcity. Authentication of this species was needed to protect consumers and conservation measures and to find out the alternative source. Deoxyribose nucleic acid (DNA) extracted from fifteen samples of six species belonging to different localities, were used as templates. Four candidate barcodes were amplified by polymerase chain reaction and sequence analysis was executed by Z-Big Dye Terminator Cycle Sequencing v.3. Sequenced products were analyzed on automated applied biosystems 3730XI analyzer. Identification was performed by using molecular evolutionary genetics analysis 5 software (version 5.1). The amplification efficiency of all DNA barcodes [megakaryocyte-associated tyrosine kinase (matK), ribulose-bisphosphate carboxylase (rbcL), photosystem II protein D1- stuctural RNA- His tRNA (psbA-trnH) and internal transcribed spacer region (ITS)] was 100 %. Here, the highest interspecific divergence provided by ITS is 11.87 % and intraspecific by psbA-trnH being 10.22 % as compared to matK (5.04 %) and rbcL (0.99 %). ITS region proves robust molecular marker for differing the S. chirayita from its related adulterant species. All barcoding regions indicate that S. chirayita and S. minor both are more closely related than other Swertia species. Findings showed that DNA barcoding is an efficient tool for identification and authentication of S. chirayita. Use of S. minor as substitute to S. chirayita can be advocated.
To understand flower morphological evolution in Gentianaceae-Swertiinae, we studied generic relationships using trnL intron, mat K, and nuclear ITS sequences of a total of 13 genera and 59 species of the subtribe. The phylogenetic incongruencies between the chloroplast and nuclear genes are likely to be the result of long branch attraction. The East Asian Megacodon and Latouchea and the eastern North American Bartonia and Obolaria were determined as the most basal genera, and several well-supported subgroups were revealed. Swertia, Lomatogonium, and Gentianella s. l. were highly polyphyletic and the position of Veratrilla and several species was ambiguous. The main flower types found in Swertiinae can be transformed into each other by simple developmental variation in proportion. This apparently happened several times during the evolution of Swertiinae and, in conjunction with other homoplastic characters, explains the difficulty of recognizing generic limits and the mosaic pattern of character distribution. Phylogenetic relationships, extant distribution ranges, and a preliminary molecular clock approach led to the hypothesis that the last common ancestor of the Swertiinae lived approximately 15 mya, and that an exchange of lineages between East Asia and North America happened frequently from the time of origin until only recently.
— We studied sequence variation in 16S rDNA in 204 individuals from 37 populations of the land snail Candidula unifasciata (Poiret 1801) across the core species range in France, Switzerland, and Germany. Phylogeographic, nested clade, and coalescence analyses were used to elucidate the species evolutionary history. The study revealed the presence of two major evolutionary lineages that evolved in separate refuges in southeast France as result of previous fragmentation during the Pleistocene. Applying a recent extension of the nested clade analysis (Templeton 2001), we inferred that range expansions along river valleys in independent corridors to the north led eventually to a secondary contact zone of the major clades around the Geneva Basin. There is evidence supporting the idea that the formation of the secondary contact zone and the colonization of Germany might be postglacial events. The phylogeographic history inferred for C. unifasciata differs from general biogeographic patterns of postglacial colonization previously identified for other taxa, and it might represent a common model for species with restricted dispersal.