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Gentianella macrosperma, a new species of Gentianella (Gentianaceae) from Xinjiang, China
expand article infoHai-Feng Cao, Ji-Dong Ya§, Qiao-Rong Zhang§, Xiao-Jian Hu§, Zhi-Rong Zhang§, Xin-Hua Liu|, Yong-Cheng Zhang, Ai-Ting Zhang#, Wen-Bin Yu¤«
‡ Shanghai University of Traditional Chinese Medicine, Shanghai, China
§ Kunming Institute of Botany, Chinese Academy of Sciences, Kunming, China
| Xinjiang Institute of Ecology and Geography, Chinese Academy of Sciences, Xinyuan, China
¶ Forestry Bureau of Xinyuan County, Xinyuan, China
# Xinjiang Agricultural Broadcasting and Television School, Xinyuan, China
¤ Center of Conservation Biology, Core Botanical Gardens, Mengla, China
« Xishuangbanna Tropical Botanical Garden, Mengla, China
Open Access

Abstract

Gentianella macrosperma Ma ex H.F. Cao, J.D. Ya & Q.R. Zhang, a new species of Gentianaceae from Xinjiang, Northwest China is described and illustrated. This new species is unique in having equal length of corolla lobe and corolla tube, nectaries located at the throat of the corolla tube and large seeds up to 1.6 mm in diameter. In addition, an updated identification key to the Chinese species of Gentianella is provided.

Keywords

Gentianella, ITS, matK, Morphology, Swertiinae, Taxonomy, Xinjiang

Introduction

Gentianella Moench (Gentianaceae) consists of approximately 300 species distributed from the temperate, arctic and alpine regions of the Northern Hemisphere, to South America, Australia and New Zealand (Pringle 2017). About 70% of species (ca. 200 species) occur in South America, where new species continue to be discovered (Pfanzelt et al. 2015; Pringle 2015, 2017; Pringle and Grant 2017). Molecular phylogenetic studies indicated that Gentianella was polyphyletic, and the new circumscription of Gentianella s. str. contains species with one nectary per petal lobe (von Hagen and Kadereit 2001, 2002). However, the taxonomic placement of the Asiatic species with two nectaries per corolla lobe has yet to be determined. Before the phylogenetically-based concept of Asiatic gentianellas proposed, the description of this genus published in Flora of China (Ho and Pringle 1995) remains applicable in the present context. There are 10 species of Gentianella reported from China and mainly distributed in northern China and alpine areas of southwest China mountains (Ho and Pringle 1995, Chen et al. 2011).

During the field expedition to west of Xinjiang, China, an unusual species of Gentianaceae was collected. Its corolla campanulate without plicae and fringed scale, lobed to middle of corolla, two nectaries per corolla lobe located at the corolla tube fit the main characters of Gentianella. Subsequent morphological investigation and molecular study supported this species as new to science and described here.

Materials and methods

Specimen collections of Gentianella were carefully examined, especially the relevant species, including G. holosteoides Schott & Kotschy ex N.M. Pritch., G. longicarpa (Gilli) Holub, G. sibirica (Kusn.) Holub, G. stoliczkae (Kurz ex C.B. Clarke) Holub and G. umbellata (M. Bieb.) Holub. Collections at the following herbaria (BM, FR, GH, GLM, HIMC, HNWP, JE, K, E, KFTA, KUN, MA, MPU, MW, P, PE, PEY, W, WAG) were checked on-site and via Chinese Virtual Herbarium (CVH, http://www.cvh.ac.cn/), Global Biodiversity Information Facility (GBIF, https://www.gbif.org/) and Global Plants on JSTOR (https://plants.jstor.org/). The high-resolution images of type specimen of G. sibirica (LE01043410, LE01043411, LE00050650) were obtained from curators of LE. Relevant literatures were investigated (Gillett 1957; Shishkin and Bobrov 1967; Omer et al. 1988; Ho and Pringle 1995; Omer 1995; Struwe et al. 2002; Aitken 2007; Chen et al. 2011; Mohd et al. 2018). Line drawings, description and most of photographs were based on the latest collections (J.D. Ya et al. 17CS16327), except that the images of seeds were from the type specimen (Shun-Li Chen Tianyi281, PE00029466). The conservation status of the new species was evaluated according to the guidelines of the IUCN Red List Categories and Criteria (IUCN 2017)

Fresh leaves of this new species were dried immediately by using silica gel for DNA extraction. Genomic DNA extraction, amplification and DNA sequencing of ITS and the plastid matK followed the protocol described by Xi et al. (2014) and sequences of relevant species were downloaded from GenBank (Appendix 1).

The molecular phylogenetic tree of 88 species representing 13 genera of Gentianaceae was reconstructed using Bayesian Inference (BI) and Maximum Likelihood (ML). Chelonanthus alatus (Aubl.) Pulle (Gentianaceae: Helieae) was chosen as outgroup (Figure 1). ITS and matK datasets were combined for analysis. BI analysis was performed using MrBayes 3.26 (Ronquist and Huelsenbeck 2003). Markov Chain Monte Carlo (MCMC) analysis was performed using MrBayes for 10,000,000 generations for the combined dataset, with two simultaneous runs, with each run comprising four incrementally heated chains. BI analysis was started with a random tree and sampled every 1000 generations. The combined dataset was partitioned and the best-fit DNA substitution model for two DNA regions using Bayesian Information Criterion (BIC) was estimated using jModeltest 2 (Darriba et al. 2012). ML analysis was conducted with RAxML 8.2.10 (Stamatakis et al. 2008) using the GTR substitution model with gamma-distributed rate heterogeneity amongst sites and the proportion of invariable sites estimated from the data. Support values for nodes/clades were estimated from 1000 bootstrap replicates.

Figure 1. 

The major-rule consensus tree of ML analysis based on the total dataset, including ITS and matK. ML bootstrap values and BI posterior probabilities are shown on branches.

Results

The ITS matrix was 689 bp in length including 376 variable sites and 266 parsimony-informative sites and the matK matrix was 821 bp in length including 286 variable sites and 198 parsimony-informative sites. The best-fit BIC model of ITS and matK datasets was SYM+G and TVM+G, respectively. The major-rule consensus tree of both BI and ML analyses with support values is shown in Figure 1.

Phylogenetic analyses using ML and BI methods identified that Gentianella, Swertia L. and other genera in subtribe Swertiinae are not monophyletic, which shows a similar conclusion as previous studies (von Hagen and Kadereit 2001, 2002; Xi et al. 2014). Current new species and 44 other Gentianella species were strongly supported as monophyletic (BI PP = 1.00, ML BS = 93; Figure 1). G. arenaria (Maxim.) T.N. Ho, G. angustiflora H. Smith, G. azurea (Bunge) Holub, G. gentianoides (Franch.) H. Smith and G. moorcroftiana (Wall. ex G. Don) A. Shaw formed different clades with Comastoma Toyok., Lomatogonium A. Braun, Swertia and other genera in Swertiinae.

Phylogenetic analyses showed that this new species and G. holosteoides formed a clade (BI PP = 0.93), then sister to the clade including G. aurea (L.) H. Smith, G. umbellata and G. longicarpa (Figure 1). Three samples of G. stoliczkae were located at most basal of the new species clade (BI PP = 1.00, ML BS = 79).

Taxonomic treatment

Gentianella macrosperma Ma ex H.F. Cao, J.D. Ya & Q.R. Zhang, sp. nov.

Figures 2, 3

Diagnosis

Resembles G. holosteoides, G. longicarpa, G. sibirica, G. stoliczkae and G. umbellata, but differs from them by having even flower size, corolla white, corolla lobe as long as corolla tube, nectaries located close to the throat of the corolla tube and larger seeds.

Type

CHINA. Xinjiang: Ili Kazak Autonomous Prefecture, Gongliu County, Ji’ergelang Township, Qiaxi Village, on the mountain ridge in the forest, 1780 m elev., 6 September 1956, Shun-Li Chen Tianyi281 (holotype: PE00029466!; isotype: PE00029453!, PE00029471!).

Description

Herbs, annual. Roots slender, yellow. Stems 30–40 cm, erect, subquadrangular, glabrous, yellowish-green, 2.0–2.5 mm in diameter; branched from the base in axils of each node, more slender, suberect or slightly ascending. Leaves opposite, basal leaves not rosette and withered at anthesis, petiole conspicuous, 7–10 mm long, leaves oblong-spatulate, 14–17 × 2–6 mm, base tapering into petiole, margin entire, apex rounded, veins 3–5, raised abaxially and slightly sunken adaxially; lower cauline leaves obovate-spatulate or rounded-spatulate, petiole 10–18 mm long, leaf blades with petiole 18–31 × 10–11 mm, both surfaces glabrous, base tapering into conspicuous petiole, margin entire, apex rounded, veins 5–7 raised abaxially and slightly sunken adaxially; middle leaves on primary stem elliptic, ovate-elliptic, 25–38 × 10–15 mm, base rounded or truncate, inconspicuously short or subsessile, both surfaces glabrous, margin entire, apex rounded, veins 5–9, raised abaxially and slightly sunken adaxially; upper stem leaves ovate-elliptic to ovate, 15–25 × 7–12 mm, with terminal two pairs of leaves nearly in whorls, both surfaces glabrous, base rounded, sessile, margin entire, apex acute, veins 3–5, raised abaxially and sunken adaxially; lateral branches leaves smaller, 10–15 × 4–7 mm. Cymes terminal and axillary, 3–4 flowers per leaf axil, terminal inflorescence 8–10 flowers, dense, inflorescence flowering at different times, pedicel variable in length and up to 36 mm. Flowers 4-merous (rarely 5-merous), all flowers almost the same size (terminal corolla as long as or slightly longer than others), rotating arrangement. Calyx 3.5–4.5 mm long, slightly shorter than corolla or as long as corolla, divided almost to the base, calyx tube 0.7–0.8 mm long, membranous, lobes green, distinctly unequal, 2 slightly larger, oblanceolate to linear-oblanceolate, 3.0–3.5 × 0.7–1.0 mm, 2 (–3) slightly smaller, linear, 2.3–3.0 × 0.4–0.5 mm, apex acute or acuminate, margin scabrous, midvein raised abaxially, sinus obtuse. Corolla white, campanulate, 4.0–4.5(5.0) mm long; corolla tube 2.1–2.4 mm long; lobes ovate, with light brown fine longitudinal veins, 2.2–2.5 × 1.5–1.8 mm, apex obtuse and mucronate, margin entire. Nectaries 8(–10), green, oblong, naked and indistinct, two nectaries per corolla lobe located very close to the throat of the corolla tube, ca. 0.2 mm from the top of corolla tube. Stamens inserted at middle of corolla tube, filaments white, linear, 1.1–1.4 mm long, anthers blue, rectangular, 0.2–0.3 mm long; ovary elliptic, ca. 2.0 mm long. Style short, linear, 0.4–0.5 mm long, stigma small, 2-lobed. Gynophore short, 0.2–0.3 mm long. Capsule elliptic, a concavity sometimes present in the centre, 2.5–4.0 mm long, usually with 2–8 seeds each capsule. Seeds brown, glossy, flat-ellipsoid, 1.2–1.6 × 0.5–0.9 mm, seed coat wrinkled-reticulate (smooth when immature).

Figure 2. 

Gentianella macrosperma, sp. nov. A plant B flower, top views C–D show opened corollas, 4- and 5-merous, respectively E flower, showing the length of calyx and corolla subequal F calyx. showing 4-merous G calyx, showing 5-merous H capsule I seeds. Drawn by R.M. Zhang. H and I from the isotype S.L. Chen Tianyi281 (PE00029471), others from the paratype J.D. Ya, Q.R. Zhang & X.J. Hu 17CS16327 (KUN1443565). Scale bars: 2 cm (A); 5 mm (B); 2 mm (C–H); 0.5 mm (I).

Figure 3. 

Gentianella macrosperma, sp. nov. A plant in nature habitat B flowers and inflorescence C flowers, showing pedicels and upper leaves D–E front view and side view of corolla, showing nectaries located close to the throat of the corolla tube F middle cauline leaf, abaxial view, showing veins G plants specimen (from KUN1443554) H opened corolla (5-merous) showing ovary I calyx J seed, front view (left and middle) and side view (right) (from S.L. Chen Tianyi281 (PE00029471)). I, H from the paratype J.D. Ya, Q.R. Zhang & X.J. Hu 17CS16327 (KUN1443565). Scale bars: 5 cm (A, G); 2 cm (B); 2 mm (C–E, I, H); 1 mm (J).

Phenology

Flowering and fruiting from June to September.

Distribution and habitat

G. macrosperma is distributed in Gongliu county and Xinyuan county, west of Xinjiang, China. It grows in thickets on the slope or on the mountain ridge in the forest of Picea schrenkiana Fisch. & Mey. at an elevation of 1729–1780 m.

Etymology

The specific epithet “macrosperma” refers to the larger seeds of this new species.

Vernacular name

Chinese mandarin: da zi jia long dan (大籽假龙胆)

Conservation status

Currently only known from three localities in west of Xinjiang, therefore considered to be Vulnerable (VU D2) (IUCN 2017).

Additional specimens examined (paratypes)

CHINA. Xinjiang: Ili Kazak Autonomous Prefecture, Xinyuan County, on the road from Xinyuan County to the gold mine, 43°16'06.45"N, 83°17'42.90"E, 1729 m elev., 1 July 2017, J.D. Ya, Q.R. Zhang & X.J. Hu 17CS16327 (KUN1443565!, KUN1443566!, KUN1443554!); Ili Kazak Autonomous Prefecture, Gongliu County, Mohuer Township, Damohe Village, 8 August 1976, Shu-Run Liu s.n. (HIMC0026063!, HIMC0026064!. the sheet 0026064 presents a mixture of Swertia dichotoma Linn. which was labelled as “A” and G. macrosperma labelled as “B”)

Discussion

It was Prof. Yu-Quan Ma (also as Yu Chuan Ma), a specialist of Gentianaceae, who first recognised this plant as a distinct new species and inscribed the name “Gentianella macrosperma Ma” on the specimen kept at PE. Later the same year, he proposed another name “Gentianella procumbens Ma” to the same collections, corresponding to its procumbent stems. However, both names were never published. Based on field observation and specimen examination, procumbent stems occurred occasionally in some individuals, the character of larger seeds being easily distinguished from other Gentianella species.

In all the known Chinese species of Gentianella, the length of corolla lobes is shorter than that of the corolla tube and nectaries which are located at the base or middle of the corolla tube. The same length of corolla lobes and corolla tube and nectaries positioned at the throat of the corolla tube make G. macrosperma a distinctive species amongst them. Its large seeds up to 1.6 mm in diameter are perhaps unique amongst the Asiatic species of Gentianella.

G. macrosperma is similar in size and shape of the corolla lobe to G. sibirica and G. longicarpa, but further differs from them both in the lack of rosette basal leaves, predominant 4-merous flowers and smaller corolla, no more than 5 mm long, except the corolla lobed to the middle, nectaries position and seeds size. Gentianella longicarpa, which is endemic to Afghanistan, is also distinct from G. macrosperma in its light-pink, pale blue or lilac-violet flower and larger corolla up to 8 mm long and all calyx lobes are shorter than the corolla tube. G. macrosperma is similar in habit and inflorescences to G. umbellata and G. stoliczkae. The flower of G. umbellate is larger than those of G. macrosperma and, although the size of the corolla lobe in the two species overlaps, the corolla lobe is much shorter than the corolla tube in G. umbellate. In G. stoliczkae, flowers are in densely clustered cymes, the corolla are generally much larger up to 20 mm long with various colours from purple, pink, pale blue to yellow and the capsule has a short gynophore ca. 1–2 mm long.

The molecular evidence shows that G. macrosperma has the closest relationship with G. holosteoides which is native to Turkey and Pakistan and they also share similar floral whorls and basal leaves shape, but plants of G. holosteoides are smaller in stature, no more than 5 (7) cm height; it further differs from G. macrosperma in its smaller basal leaves, larger flowers with corolla lobes shorter than corolla tube, nectaries position at corolla base and smaller, numerous seeds. A detailed morphological comparison is given in Table 1.

Morphological comparison between Gentianella macrosperma and related species.

G. macrosperma G. holosteoides G. longicarpa G. sibirica G. stoliczkae G. umbellata
Plant height (cm) 12–40 up to 5 9–22 (1–)10–20(–30) 10–45(–60) (4–)10–35
Basal leaves (mm) not rosulate, obovate-spathulate 14–17 × 2–6 rosulate, spathulate-ovate or lanceolate, 3–5 × 1–3 rosulate, spathulate, oblong-obovate, 7–16 × 3–8 rosulate, oblong-obovate, 6–20 × 2–6 rosulate, ovate-lanceolate to ovate, 10–35 × 6–20 rosulate, spathulate, obovate-lanceolate, 8–25 × 5–12
Cauline leaves (mm) ovate to ovaloid, apex rounded, the uppermost sometimes acute, 15–38 × (7–)10–15 lanceolate-oblanceolate or elliptic, apex acute, 5–15 × 2–6 ovate-oblong, ovate or ovate-lanceolate, apex obtuse, the uppermost acute, 8–26 × 4–9 ovate-oblong, ovate-lanceolate, apex acute, 6–20(–35) × 3–9 oblong-lanceolate, lanceolate to ovate-lanceolate, apex acute, (20–)25–40(–50) × (2–)10–15 oblong-ovate, oblong-lanceolate, apex acute, 8–25 × 4–18
Calyx length (mm) 3.5–4.5 4–8 4–5 3–6 8–11 4–10
Floral whorls 4(5)–merous 4(5)–merous 5-merous 5(4)-merous 5-merous 5-merous
Flower size almost all of the same size variable in size, terminal ones 1–2 × larger than others variable in size, terminal ones 1–1.5 × larger than others variable in size, terminal ones 1–2 × larger than others variable in size, terminal ones 2–3 × larger than others variable in size, terminal ones 2–3 × larger than others
Corolla colour white pale blue to blue pale blue, light-pink, or lilac-violet predominantly pink, yellowish or whitish, rarely pale blue. purple, pink, pale blue or yellowish pale azure, purple, pink, yellowish or mixture of these, rarely white
Corolla shape campanulate tubular to campanulate-tubular tubular to campanulate-tubular tubular or tubular-infundibular tubular to campanulate-tubular tubular to campanulate-tubular
Corolla length (mm) 4.0–4.5(–5.0) 6–12 (5–) 6–8 (5–) 6–7 (–10) 7–20 (5–) 8–11 (–15)
Corolla lobes 2.0 mm long, the same length as corolla tube 1.5–3.0 mm long, much shorter than corolla tube 2–3 mm long, shorter than corolla tube ca. 2 mm long, much shorter than corolla tube 3–7 mm long, much shorter than corolla tube 2–3(4) mm long, much shorter than corolla tube
Nectaries 8(10), at top of corolla tube 8(10), at basal part of corolla tube 10, at basal part of corolla tube 8-10, at basal part of corolla tube 10, at basal part of corolla tube 10, at basal part of corolla tube
Stamens 1.1–1.4 mm 2–4 mm ca. 7 mm 1–5 mm
Anthers blue, 0.2–0.3 mm blue, 0.5–0.7 mm yellow, 1.0–1.2 mm
Gynophore 0.2–0.3 mm subsessile sessile subsessile 1.5–2.2 mm sessile
Seeds 2–8 per capsule, 1.2–1.6 mm in diameter numerous per capsule, ca. 0.8–1.0 mm in diameter numerous per capsule, 0.2–0.3 mm in diameter numerous per capsule, 0.1–0.2 mm in diameter numerous per capsule, ca. 0.8 mm in diameter numerous per capsule, 0.2–0.3 mm in diameter

Von Hagen and Kadereit (2001) proposed Gentianella s. str. to only include species with one nectary per petal lobe, however, G. umbellata and G. stoliczkae represented in their study are both binectariate species. Current molecular analyses also shows the binectariate G. macrosperma clustered into von Hagen and Kadereit’s Gentianella s. str. A careful selection of species across wider geographic regions of this genus and data from more nuclear and chloroplast sequences may clarify the generic circumscription in Gentianella.

Key to species of Gentianella in China

The following key is based on Flora of China (Ho and Pringle 1995), Flora of the U.S.S.R. (Shishkin and Bobrov 1967) and other literature (Omer et al. 1988; Aitken 2007; Chen et al. 2011). It includes 11 species of Gentianella in China.

1 Corolla lobes fimbriate at base G. acuta
Corolla lobes glabrous at base 2
2 Nectaries above the middle of corolla tube 3
Nectaries at the base of corolla tube 4
3 Plant 12–40 cm tall, nectaries close to the throat of corolla tube, seeds 1.2–1.6 mm in diameter G. macrosperma
Plant 1–4 cm tall, nectaries just above the middle of corolla tube, seeds 0.7–0.8 mm in diameter G. pygmaea
4 Margin and midvein of calyx lobe blackish G. azurea
Calyx not as above 5
5 Stem densely purple pilose G. gentianoides
Stem glabrous (sometimes sparsely pilose in G. moorcroftiana) 6
6 Flowers often angled, corolla tube 3–4 times longer than lobe G. angustiflora
Flowers not angled, corolla tube 1–3 time(s) longer than lobe 7
7 Corolla lobes apically obtuse or round 8
Corolla lobes apically mucronate 9
8 Flowers 5-merous, stem leaf blades linear G. moorcroftiana
Flowers 4-merous, stem leaf blades spatulate to oblong-spatulate G. arenaria
9 Corolla lobes densely papillate outside G. anomala
Corolla lobes glabrous outside 10
10 Corolla 7–20 mm long, terminal ones ca. 20 mm, lobes 3–7 mm G. stoliczkae
Corolla 4–10 mm long, terminal ones up to 10 mm, lobes ca. 2 mm G. sibirica

Acknowledgements

We are grateful to three anonymous reviewers and Jie Cai for their critical comments to improve the manuscript; to En Zhou of Shanghai University of Traditional Chinese Medicine for language revision; to Fu-Lin Li and Jian-Zhong Pang from Yeguolin improvement station of Xinyuan for their kind assistance in the field; to Lian-Yi Li of Kunming Institute of Botany, Chinese Academy of Sciences (CAS) for image processing; to Irina Illarionova and Larisa Raenko of LE, Wei Zhao and Yang-Jun Lai of Institute of Botany, CAS for specimen images access; to Rong-Mei Zhang for the line drawing. We appreciate curators of the cited herbaria and the website managers of CVH, GBIF, JSTOR for the online images access. This study was financially supported by CAS’s Large-scale Scientific Facilities (Grant No.: 2017-LSF-GBOWS-02) and Basic Research Project of the Ministry of Science and Technology of China (Grant No.: 2013FY112600).

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Appendix

Appendix 1.Download as CSV 

Samples for phylogenetic analysis using matK and ITS sequences with voucher information, GenBank accession number.

Species Voucher specimen (Herbarium/No.) Locality matK ITS
OUTGROUP
Chelonanthus alatus (Aubl.) Pulle Maas 9316 (U) French Guiana KX904551 KX904610
GENTIANEAE GROUP
Crawfurdia speciosa Wall. KEKE 1244 (K) N/A AJ010512/AJ011441 AJ294586/AJ294646
Gentiana crassicaulis Duthie ex Burkill xuechy090107 (KUN) China KC861277 KC861348
Gentiana dahurica Fisch. xuechy0076 (KUN) China KC861279 KC861350
Gentiana frigida Haenke N/A Germany (Schachen Bot. Garden), cultivated AJ388166/AJ388236 AJ294588/AJ294648
Gentiana straminea Maxim. xuechy0065 (KUN) China KC861282 KC861353
SWERTIINAE GROUP
Comastoma cyananthiflorum (Franch.) Holub XHC120021 (KUN) China KC861250 KC861320
Comastoma cyananthiflorum (Franch.) Holub CEE-88 (E 00025334) China AJ406324/AJ406353 AJ294585/AJ294645
Comastoma jigzhiense T.N. Ho & J.Q. Liu Chensl0423 (KUN) China KC861231 KC861300
Comastoma polycladum (Diels & Gilg) T.N. Ho xuechy090036 (KUN) China KC861275 KC861346
Comastoma pulmonarium (Turcz.) Toyok. GLM-081307 (KUN) China KC861238 KC861306
Frasera albicaulis Griseb. K. Gutsche 20 (MJG) N/A AJ406325/AJ406354 AJ294587/AJ294647
Gentianella amarella (L.) Börner W.J. Schrenk (FR) N/A AJ406326/AJ406355 AJ294591/AJ294651
Gentianella angustiflora H. Smith Edinburgh Makalu Expedition 430 (E 00025322) Nepal AJ406327/AJ406356 AJ294592/AJ294652
Gentianella antipoda (Kirk) T.N. Ho & S.W. Liu CHR 510015 New Zealand AY136500
Gentianella arenaria (Maxim.) T.N. Ho T.N. Ho et al. 435 (E 00025341) N/A AJ406328 AJ294593/AJ294653
Gentianella aspera (Hegetschw.) Dostál ex Skalický, Chrtek & Gill K. Gutsche 45 (MJG) N/A AJ010517/AJ011446 AJ294594/AJ294654
Gentianella astonii (Petrie) T.N. Ho & S.W. Liu CHR 509942 New Zealand AY136494
Gentianella aurea (L.) H. Smith H. Smith 4131 (E 00025348) N/A AJ406329/AJ406357 AJ294595/AJ294655
Gentianella auriculata (Pall.) J.M. Gillett C. Tpyrgeba (K) N/A AJ406330/AJ406358 AJ294596/AJ294656
Gentianella austriaca (A. Kern. & Jos.Kern.) Holub N/A (MJG) Germany (Schachen Bot. Garden), cultivated AJ294597/AJ294657
Gentianella azurea (Bunge) Holub xuechy090033 (KUN) China KC861284 KC861355
Gentianella azurea (Bunge) Holub T.N. Ho, B. Bartholomew, M. Gilbert 1312 (E 00025339) China AJ406331/AJ406359 AJ294598/AJ294658
Gentianella azurea (Bunge) Holub Yangyp-Q-0255 (KUN) China MN067526* MK416127*
Gentianella bellidifolia (Hook.f.) Holub 19932974 Scotland (Edinburgh Bot. Garden), cultivated AJ388162/AJ388232 AJ294599/AJ294659
Gentianella bohemica Skalický 015 Czech Republic AJ580570
Gentianella canosoi G.L. Nesom & B.L. Turner S. Gonzales, S. Acevedo 2033 (TEX) N/A AJ406332/AJ406360 AJ294600/AJ294660
Gentianella caucasea (Lodd. ex Sims) Holub J. C. Archibald 8208 (E 00025347) N/A AJ294601/AJ294661
Gentianella cerastioides (Kunth) Fabris R. Greissl (MJG) N/A AJ010518/AJ011447 AJ294602/AJ294662
Gentianella cernua (Kunth) Fabris C.Viteri 4410 (MO) N/A AJ294603/AJ294663
Gentianella cosmantha (Griseb.) J.S. Pringle J.G. Haukes, J.P.Hjirting, K. Rahn 3569 (L 424359) N/A AJ406333/AJ406361 AJ294604/AJ294664
Gentianella diemensis (Griseb.) J.H. Willis H. Hurka (MJG) N/A AJ295332/AJ295333 AJ294605/AJ294665
Gentianella engadinensis (Wettst.) Holub Ge002 Switzerland AJ580559
Gentianella fastigiata (Benth.) Fabris K. Gutsche (MJG) N/A AJ294606/AJ294666
Gentianella florida (Griseb.) Holub R. Ehrich 444 (MJG) N/A AJ406334/AJ406362 AJ294607/AJ294667
Gentianella foliosa (Kunth) Fabris 1994-508 England (Kew Bot. Garden), cultivated AJ294608/AJ294668
Gentianella gentianoides (Franch.) H. Smith xuechy090065 (KUN) China KC861285 KC861356
Gentianella gentianoides (Franch.) H. Smith xuechy090094 (KUN) China KC861286 KC861357
Gentianella germanica (Willd.) E.F. Warburg 20 Germany AJ580562
Gentianella germanica (Willd.) E.F. Warburg J.W. Kadereit (MJG) N/A AJ406335/AJ406363 AJ294609/AJ294669
Gentianella hirculus (Griseb.) Fabris J.L. Clarke 1787 (QCNE) N/A AJ294610/AJ294670
Gentianella holosteoides Schott & Kotschy ex N.M. Pritch. Southhampton University 179 (K) N/A AJ294611/AJ294671
Gentianella macrosperma Ma ex H.F. Cao, J.D. Ya & Q.R. Zhang 17CS16327 (KUN) China MN067523* MK416132*
Gentianella lineata (Kirk) T.N. Ho & S.W. Liu CHR 509866 New Zealand AY136503
Gentianella longicarpa (Gilli) Holub D. Podlech 12436 (M) N/A AJ294612/AJ294672
Gentianella magellanica (Gaudich.) Fabris K. Kubitzki, T. Feuerer 99-10 (MJG) N/A AJ406336/AJ406364 AJ294613/AJ294673
Gentianella microcalyx (Lemmon) J. M. Gillett E. Joyal, J. Enrique 1853 (TEX) N/A AJ406337/AJ406365 AJ294614/AJ294674
Gentianella montana (G. Forst.) Holub CHR 509944 New Zealand AY136491
Gentianella moorcroftiana (Wall. ex G. Don) A. Shaw R. McBeath 2093 (E 00025318) N/A AJ406338/AJ406366 AJ294615/AJ294675
Gentianella narcissoides (Gilg) T.N. Ho & S.W. Liu L. Naessany 14 (MJG) N/A AJ294616/AJ294676
Gentianella patula (Kirk) Holub 19932978 Scotland (Edinburgh Bot. Garden), cultivated AJ406339/AJ406367 AJ294617/AJ294677
Gentianella peruviana (Griseb.) Fabris 19950534 Scotland (Edinburgh Bot. Garden), cultivated AJ388163/AJ388233 AJ294618/AJ294678
Gentianella propinqua (Richardson) J.M. Gillett G. Halliday A 333/75 (E 00025300) North America AJ406340/AJ406368 AJ294619/AJ294679
Gentianella quinquefolia (L.) Small Bozeman, Ramseur, Radford 45200 (E 00025241) North America AJ406341/AJ406369 AJ294620/AJ294680
Gentianella quinquefolia (L.) Small D. Pittillo 12106 (WCUH) America EU812469
Gentianella rapunculoides (Willd. ex Schult.) J.S. Pringle R. Greissl 616 (MJG) N/A AJ294621/AJ294681
Gentianella ruizii (Griseb.) Holub Weigend, Weigend 2000/386 (NY) N/A AJ406342/AJ406370 AJ294622/AJ294682
Gentianella rupicola (Kunth) Holub 199930516 Scotland (Edinburgh Bot. Garden), cultivated AJ294623/AJ294683
Gentianella saxosa (G. Forst.) Holub Gutsche (MJG) N/A AJ406343/AJ406371
Gentianella splendens (Gilg) Fabris J.L. Clarke 1855 (QCNE) N/A AJ295336/AJ295337 AJ294624/AJ294684
Gentianella stoliczkae (Kurz ex C.B. Clarke) Holub LiuJQ0028 (KUN) China MN067524* MK416130*
Gentianella stoliczkae (Kurz ex C.B. Clarke) Holub LiuJQ0071 (KUN) China MN067525* MK416131*
Gentianella stoliczkae (Kurz ex C.B. Clarke) Holub O. Anders 8178 (M 50043) N/A AJ406344/AJ406372 AJ294625/AJ294685
Gentianella sulphurea (Gilg) Fabris J.L. Clarke 1833 (QCNE) N/A AJ294626/AJ294686
Gentianella thyrsoidea (Hook. f.) Fabris D.N. Smith, F. Escalona 10134 (MO) N/A AJ294627/AJ294687
Gentianella tristicha (Gilg) Fabris ex T.N. Ho & S.W. Liu D.N. Smith, F. Escalona 10125 (MO) N/A AJ294628/AJ294688
Gentianella umbellata (M. Bieb.) Holub K91-G3 Georgia Z48102/Z48132
Gentianella wislizenii (Engelm.) J.M. Gillett M. Lavin 4947 (TEX) N/A AJ294630/AJ294690
Gentianopsis barbata (Froel.) Ma xuechy090085 (KUN) China KC861287 KC861358
Gentianopsis crinita (Froel.) Ma N/A (MJG) Germany (Mainz Bot. Garden), cultivated AJ406345/AJ406373 AJ294631/AJ294691
Halenia elliptica D. Don GLM-081543 (KUN) China KC861242 KC861310
Halenia palmeri A. Gray K.B.v. Hagen 98/41 (MJG) N/A - AJ294632/AJ294692
Jaeschkea oligosperma (Griseb.) Knobl. R. McBeath 2300 (E 00025275) N/A AJ388171/AJ388241 AJ294633/AJ294693
Lomatogonium bellum (Hemsl.) H. Smith GLM-06075 (KUN) China KC861237 KC861305
Lomatogonium carinthiacum (Wulfen) Rchb. V. Zuev 6649 (BR) N/A AJ406346/AJ406374 AJ294634/AJ294694
Lomatogonium forrestii (I.B. Balfour) Fernald XHC120061 (KUN) China KC861261 KC861332
Lomatogonium gamosepalum (Burkill) H. Smith GLM-081372 (KUN 1272996) China KC861241 KC861309
Lomatogonium oreocharis (Diels) C. Marquand CLD-90 1106 (K) N/A AJ388174/AJ388244 AJ294635/AJ294695
Megacodon stylophorus (C.B. Clarke) H. Smith GLM-081957 (KUN) China KC861245 KC861313
Megacodon stylophorus (C.B. Clarke) H. Smith Kuming, Edinburgh, Gothenburgh Exp. 1378 (E 00025279) China AJ388177/AJ388247 AJ294636/AJ294696
Swertia bifolia Batalin Chensl0388 (KUN) China KC861229 KC861298
Swertia bimaculata (Sieb. & Zucc.) Hook. f. & Thomson ex C.B. Clarke XHC120026 (KUN) China KC861264 KC861335
Swertia bimaculata (Sieb. & Zucc.) Hook. f. & Thomson ex C.B. Clarke XCY090050 (KUN) China JF956557 JF978820
Swertia cincta Burkill XCY090098 (KUN) China JF956561 JF978823
Swertia crassiuscula Gilg U. Hecker 1094 (MJG) N/A AJ406347/AJ406375 AJ294637/AJ294697
Swertia decora Franch. XCY090077 (KUN) China JF956567 JF978825
Swertia erythrosticta Maxim. xuechy090044 (KUN) China KC861267 KC861338
Swertia franchetiana H. Smith XHC120048 (KUN) China KC861256 KC861326
Swertia japonica (Schult.) Makino N/A (KYO) Japan (Kyoto Bot. Garden), cultivated AJ406348/AJ406376 AJ294638/AJ294698
Swertia macrosperma (C.B. Clarke) C.B. Clarke XHC120060 (KUN) China KC861260 KC861331
Swertia macrosperma (C.B. Clarke) C.B. Clarke J.H. de Haas 2765 (U 500099) N/A AJ406349/AJ406377 AJ294639/AJ294699
Swertia nervosa (G. Don) Wall. ex C.B. Clarke XHC120053 (KUN) China KC861258 KC861328
Swertia patens Burkill 09CS1123 (KIB) China KC861233 KC861302
Swertia perennis L. K.B. Hungerer (MJG) N/A AJ294640/AJ294700
Swertia punicea Hemsl. 19943574 Scotland (Edinburgh Bot. Garden), cultivated AJ406350/AJ406378 AJ294641/AJ294701
Swertia racemosa (Wall. ex Griseb.) C.B. Clarke J.H. de Haas 2725 (U 500131) N/A AJ406351/AJ406379 AJ294642/AJ294702
Swertia volkensii Gilg U. Hecker 1093 (MJG) N/A AJ406352/AJ406380 AJ294643/AJ294703
Swertia yunnanensis Burkill XCY090089 (KUN) China JF956585 JF978836
Veratrilla baillonii Franch. Kuming, Edinburgh, Gothenburgh Exp. 1326 (E 00025273) China AJ388196/AJ388266 AJ294644/AJ294704