Research Article |
Corresponding author: Ming-Tai An ( gdanmingtai@126.com ) Academic editor: Rafael Felipe Almeida
© 2024 Tian-Rou Wu, Jian Xu, Ming-Tai An, Jiang-Hong Yu, Feng Liu, Zheng-Ren Chen.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Wu T-R, Xu J, An M-T, Yu J-H, Liu F, Chen Z-R (2024) Hypericum liboense (Hypericaceae), a new species from Guizhou, China. PhytoKeys 237: 37-49. https://doi.org/10.3897/phytokeys.237.110482
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Hypericum liboense M.T.An & T.R.Wu, sp. nov. (Hypericaceae) is a newly described species found in the Maolan National Nature Reserve of Guizhou Province, where it grows in rocky habitats without soil on karst mountain tops. In this study, key morphological characters were compared between the new species and the other known Hypericum species of Hypericaceae. DNA sequences were extracted from the leaves of the new species, with nuclear gene sequences (ITS) generated to reconstruct phylogenetic trees and describe its phylogenetic position in relation to other species of Hypericum. Our results show that the proposed new species has the typical characteristics of the genus Hypericum in morphology being similar to Hypericum monogynum, but differing in its sessile and semi-clasped leaves, long elliptical to long circular leaf blades, thickly papery to thinly leathery, with entire and wavy leaf margins. The abaxial side of the leaves is covered with white powder, giving them a grey-white appearance. The main lateral veins of the leaves are 8–15-paired, and the midvein on both sides is convex. The main lateral veins and midvein branch are conspicuous, with tertiary venation forming a network on the leaf surface and appearing prominently sunken. The inflorescences are 1–3-flowered, with a large calyx and conspicuous veins. The molecular phylogenetic analysis (PP = 1.00) provided substantial evidence for the proposition of H. liboense as a new species within Hypericum. Morphological and molecular evidence is presented, corroborating the proposition of the new species, including a comprehensive account of the distinctive morphological attributes of H. liboense, along with its key distinguishing features from similar species.
Molecular evidence, morphology, phylogeny, taxonomy
Hypericum L. is the largest genus of the family Hypericaceae, with approximately 470 species worldwide (
The genus Hypericum was originally classified under the family Guttiferae based on morphological studies (
The APG IV system (
In 2022, we participated in a plant survey in a karst area of the Maolan National Nature Reserve in Guizhou, China, and discovered an unusual specimen of Hypericaceae. After field investigation and collection of specimens, we conducted detailed morphological analyses and realised that the morphological characteristics of this species were similar to those of Hypericum, but there were obvious differences in the leaf and calyx from the species occurring in China. To effectively differentiate species H. liboense from others within the genus Hypericum, this study utilized phylogenetic analysis based on morphological identification and description, combined with ITS sequences. As a result, a conclusion was reached, designating it as a novel species within the realm of scientific understanding.
The ITS sequence, a highly reiterated tandem sequence in the nuclear genome, exhibits rapid changes, providing abundant variation and informative sites (
Section number | Specie number | Species | GenBank No. | Section |
---|---|---|---|---|
1 | 1 | Hypericum quartinianum A.Rich. | HE653603.1 | Campylosporus |
2 | 2 | Hypericum balearicum L. | AY555862.1 | Psorophytum |
3 | 3 | Hypericum bellum subsp. latisepalum N.Robson | HE653426.1 | Ascyreia |
4 | Hypericum calycinum L. | HE653431.1 | ||
5 | Hypericum forrestii (Chittenden) N. Robson | HE653476.1 | ||
6 | Hypericum hookerianum Wight et Arn. | KC709450.1 | ||
7 | Hypericum kouytchense Lévl. | FJ694210.1 | ||
8 | Hypericum lagarocladum N. Robson | HE662703.1 | ||
9 | Hypericum patulum Thunb. ex Murray | FJ694214.1 | ||
10 | Hypericum pseudohenryi N. Robson | KC709447.1 | ||
11 | Hypericum wilsonii N. Robson | HE653658.1 | ||
12 | Hypericum monogynum L. | HE653544.1 | ||
4 | 13 | Hypericum geminiflorum Hemsl. | HM162838.1 | Takasagoya |
5 | 14 | Hypericum androsaemum L. | KC709337.1 | Androsaemum |
15 | Hypericum grandifolium Choisy | KC709385.1 | ||
16 | Hypericum × inodorum Mill. | HE653565.1 | ||
6 | 17 | Hypericum xylosteifolium N. Robson | HE653659.1 | Lnodora |
7 | 18 | Hypericum przewalskii Maxim. | JF976672.1 | Roscyna |
8 | 19 | Hypericum bupleuroides Griseb. | HE653429.1 | Bupleuroides |
9 | 20 | Hypericum attenuatum Choisy | HE662752.1 | Hypericum |
21 | Hypericum kamtschaticum Ledeb. | HE653516.1 | ||
22 | Hypericum perforatum L. | JN811136.1 | ||
23 | Hypericum perforatum subsp. veronense (Schrank) H. Lindb. | MN036448.1 | ||
24 | Hypericum pseudopetiolatum R. Keller | AY573002.1 | ||
25 | Hypericum yezoense Maxim. | AY573004.1 | ||
10 | 26 | Hypericum concinnum Benth. | HE653442.1 | Concinna |
11 | 27 | Hypericum pseudomaculatum Bush | HE653595.1 | Graveolentia |
12 | 28 | Hypericum sampsonii Hance | HE653620.1 | Sampsonia |
13 | 29 | Hypericum elodeoides Choisy | HE653457.1 | Elodeoida |
14 | 30 | Hypericum monanthemum Hook. f. et Thoms. ex Dyer | HE653542.1 | Monanthema |
15 | 31 | Hypericum polyphyllum Boiss. & Balansa | HE662730.1 | Olympia |
16 | 32 | Hypericum cerastoides (Spach) N.Robson | AY555884.1 | Campylopus |
17 | 33 | Hypericum papillare Boiss. & Heldr. | HE653570.1 | Origanifolia |
18 | 34 | Hypericum barbatum Jacq. | FJ694192.1 | Drosocarpium |
35 | Hypericum richeri subsp. grisebachii (Boiss.) Nyman | FJ694222.1 | ||
36 | Hypericum rumeliacum Boiss. | HE653616.1 | ||
19 | 37 | Hypericum humifusum L. | HE653507.1 | Oligostema |
20 | 38 | Hypericum orientale L. | HE653565.1 | Crossophyllum |
21 | 39 | Hypericum pseudolaeve N.Robson | HE653594.1 | Hirtella |
22 | 40 | Hypericum hirsutum L. | HE653500.1 | Taeniocarpium |
41 | Hypericum pulchrum L. | FJ694219.1 | ||
23 | 42 | Hypericum empetrifolium Willd. | HE653464.1 | Coridium |
24 | 43 | Hypericum hypericoides (L.) Crantz | KC709376.1 | Myriandra |
44 | Hypericum kalmianum L. | FJ694209.1 | ||
45 | Hypericum prolificum L. | MT551029.1 | ||
25 | 46 | Hypericum canariense L. | KC709387.1 | Webbia |
26 | 47 | Hypericum vacciniifolium Hayek & Siehe | HE653656.1 | Arthrophyllum |
27 | 48 | Hypericum pallens Banks & Sol. | AY555848.1 | Triadenioides |
28 | 49 | Hypericum heterophyllum Vent. | HE653492.1 | Heterophylla |
29 | 50 | Hypericum aegypticum subsp. webbii L. | KC709380.1 | Adenotrias |
30 | 51 | Hypericum papuanum Ridl. | HE653571.1 | Humifusoideum |
31 | 52 | Hypericum reflexum L.f. | HE662747.1 | Adenosepalum |
32 | 53 | Hypericum elodes L. | FJ694200.1 | Elodes |
33 | 54 | Hypericum mexicanum L. | LT904662.1 | Brathys |
34 | 55 | Hypericum japonicum Thunb. ex Murray | HE653513.1 | Trigynobrathys |
Outgroup | Thornea calcicole Standl. & Steyerm | AY573028.1 | – |
The sequences were imported into BioEdit 7.0 for manual alignment and sorting, resulting in a refined sequence matrix, which was then exported (
During the period of 2022–2023, we conducted a field investigation on H. liboense in Maolan National Nature Reserve, Guizhou Province, including photographing its characteristics and collecting seven live specimens. The type specimen is deposited in the Tree Herbaria, College of Forestry, Guizhou University, Huaxi District, Guiyang City, Guizhou Province, China (GZAC, GZAC–LB–0001). The morphological comparison of H. liboense specimens with similar species, such as H. monogynum, was conducted by studying various materials including leaves, flowers, fructus, and branches. This comparison was primarily based on authoritative plant literature, specifically descriptions found in Flora of China (
Phylogenetic analyses indicated that the 34 included taxa of Hypericum formed a well-supported monophyletic group (Fig.
Hypericum liboense A habitat B flower (a) undulating leaf margins C flower anatomy (a) petal (b) stamens (c) pistil (d) calyx (e) anther (f) gland D H. liboense branch with flowers E stylus F veins and glandular points of calyx G leaf blade half-clasping twig H abaxial side of calyx I blade J veins and glands K anatomy of fructus (a) whole fructus (b) longitudinal section of fructus (c) cross-section of fructus
H. liboense | H. monogynum | H. kouytchense | H. patulum | |
---|---|---|---|---|
Petiole | Leaves sessile, semi-amplexicaul | Leaves sessile or brachy petiolate | Leaves petiolate, 0.5–1.5 mm long | Leaves petiolate, 0.5–2 mm long |
Leaf texture | Thickly papery to thinly leathery | Thickly papery | Thickly papery | Thickly papery |
Leaf morphology | Long elliptical to long circular | Oblanceolate or elliptic to long circular | Elliptic, lanceolate to ovate or triangular ovate | Blade lanceolate or oblong-lanceolate to ovate or oblong-ovate |
Leaf margin | Slight undulation | Flat | Flat | Flat |
Leaf lower epidermis character | Greyish white with white powder | Light green, not grey | Light green, not grey | Abaxially rather glaucous |
Midvein | Midvein raised on both sides of the leaf surface | Midvein flat on the leaf surface | Midvein flat on the leaf surface | Midvein flat on the leaf surface |
Main lateral veins | 8–15-paired | 4–6-paired | 3–4-paired | 3-paired |
Tertiary reticulation | Conspicuous, sunken on the leaf surface | Not very conspicuous | Obscure or invisible | Scarcely visible |
Inflorescence | 1–3-flowered | 1–15(–30)-flowered | 1–7(–11)-flowered | 1–15-flowered |
Anther | Yellow, glandular | Yellow to dark orange, glandular | Yellow, glandular | Yellow, glandless |
Calyx size | Elliptic or broad ovate, larger, 10–14×4–6 mm | Broad or narrowly elliptic or oblong to lanceolate or oblanceolate, smaller, 4.5–13×1.2–2 mm | Oblong-ovate to lanceolate, larger, 7–15×2.5–7 mm | Broadly ovate or broadly elliptic or subcircular to oblong-elliptic or obovate-spatulate, 5–10×3.5–7 mm |
Calyx margin | Margin entire | Margin entire | Margin entire | Margin eroded-denticulate to ciliolate with markedly hyaline margin |
Thin veins of calyx | Obvious | Obscure | Obscure | Obscure |
The results of our phylogenetic tree showed that all species of sect. Asian (including H. patulum, H. kouytchense and H. monogynum, etc) clustered together into a single clade with strong support (PP = 1.00), and the results were consistent with previous studies (
In recent years, new species of the genus Hypericaceae have been gradually discovered and reported (
China, Guizhou Province, Libo County, Maolan National Nature Reserve, elev. 947, 25°16'N, 107°57'E, 21 April 2022, Jian Xu, Mingtai An and Tianrou Wu 220421(Holotype: GZAC–LB–0001, Isotype: GZAC–LB–0002).
This species is similar to H. monogynum in terms of morphology. The main difference between the two species is that the leaves of H. liboense are sessile and semi-clasped (vs. leaves sessile or brachypetiolate). The leaves of H. liboense are long elliptical to long circular, and the edges are whole and wavy (vs. oblanceolate or elliptic to long circular, flat). H. liboense leaves are thickly papery to thinly leathery (vs. thickly papery), with a white powder on the abaxial side leading to a grey-white appearance (vs. abaxially without grey). Main lateral veins of leaves 8–15 pairs (vs. 4–6 pairs), with the midvein on both sides convex, the main lateral veins obvious branches from the midvein, the main lateral veins and tertiary vein forming an obvious network and obviously sagging (vs. tertiary vein obscure and not sunken). Inflorescences with 1–3 flowers (vs. 1–15(–30) flowers), calyx are elliptic or broad-ovate (vs. broad or narrowly elliptic or oblong to lanceolate or oblanceolate), 10–14 mm long, 4–6 mm wide (vs. 4.5–13 mm long and 1.2–2 mm wide), and veins obvious (vs. obscure) (Table
Plants Erect shrub, 0.5–1.3 m tall. Young branches reddish brown with a light white powder. Old branches dark reddish-brown or grey, cylindrical, with a lumpy rind after cracking off, and the cortex light red. Leaves opposite, sessile, with semi-clasping branchlets. The leaves are long elliptical to long circular, 4–8 cm long and 2–4 cm wide, with the middle entire part of the leaf usually the widest, the apex blunt round, with a fine cusp; leaf blade base cuneate to rounded, margin entire and slightly ruffled; thickly papery to thinly leathery, glabrous, the surface of the leaves green or dark green, the back of the leaves white and greyish-white; the main lateral veins of the leaves in 8–15 pairs, the midvein raised on both sides, and the base reddish; main lateral veins and midvein branching obviously, main lateral veins and tertiary vein forming an obvious network and concave on the leaf surface; wholly punctiform glands. Inflorescence with 1–3 flowers, emanating from the first segment of the stem; peduncle yellow-green, 1.3–3 cm long. Flowers 4–7 cm in diameter; bud ovular, apex subacute. Calyx 5, free, ovate to broadly ovate, 1–1.4 cm long, 0.4–0.6 cm wide, wholly punctiform glands, apex acute to rounded, entire margin, base light green, margin purplish red, midvein and veining obvious, and calyx enlarged in the fruit stage. Petals 5, yellow, without flush, open, triangular obovate, slightly curved, 2.8–3.5 cm long, 1.6–2.2 cm wide, approximately, margin entire, glandular. Stamens in 5 fascicles, each with 23–40 stamens, 1.3–3.4 cm long, several times the length of the petal, anthers yellow to dark orange, with glands. Ovary ovulate or sub-globular, 3–5 mm long, 2.5–5 mm wide. Style 1.3–2.2 cm long, styles partly united (style confluent almost to apex and then curved outwards into 5 splits), stigma small, lavender. Capsule broadly oval-shaped or oval-shaped and conical, 10–14 mm long and 6–10 mm wide, light green, dark brown when ripe.
Flowering from April to June; fruiting from June to September.
This species is known to be found only in Libo County, Guizhou Province, China, on the top of a mountain in a karst landscape, alt. 947 m.
The species name “liboense” refers to the origin of the type specimen, Libo County, Guizhou Province.
In the 2022–2023 period, we sampled the H. liboense population and found two more sites around the location where the species was first discovered, each with a population of approximately 20 plants. The habitat of H. liboense is mainly from the exposed rock gully area above the middle of the mountain to the top of the mountain. The soil in the plant habitat is poor, the soil layer is weak in its ability to retain water, and drought is common. At present, H. liboense is not known to be distributed in the low-altitude areas below the foot of the mountain and the middle of the mountain, so we hypothesize that the current availability of habitat for H. liboense is relatively poor and the population is relatively endangered. However, because our current investigation of the survival status and threat factors of H. liboense is not sufficiently comprehensive to provide information on the specific distribution of this population, we recommend that H. liboense be classified as “data missing” (
According to the morphological characteristics and molecular evidence of Hypericum, the findings indicate that H. liboense should be categorized within sect. Ascyreia. It is evident that H. liboense is a distinct member of Hypericum and forms a strongly supported clade (PP = 1.00) within the Hypericum phylogenetic tree. Moreover, H. liboense exhibits distinct morphological features that differentiate it from all currently accepted species in Hypericum. Therefore, it is deemed necessary to classify H. liboense as a new species.
The authors would like to thank Kun Wang, Xing-Yong Cui, Xi-Hong Yang and Hong-Fen Hu for their help with species investigation and specimen collection. Thanks to Ling-Bin Yan for his help with photographing the species.
The authors have declared that no competing interests exist.
No ethical statement was reported.
This work was supported by the "Survey and Assessment of Newly Added National Key Protected Wild Plant Resources in Guizhou Province (Second stage) (MCHC–ZD20232020)", "Research on the diversity of Corybas fanjingshanensis mycorrhizal fungi of rare and endangered orchids endemic in Guizhou (QKHJC [2023]1Y235)" and "Investigation and Monitoring Project of Maolan Large-scale Dynamic Plot of Karst Forest Eco-system in South China (2023–23)".
Tian-Rou Wu and Jian Xu completed all the work of this paper together, they contributed equally to this work and are the co-first authors of this paper. Ming-Tai An planned and guided the writing of the whole paper, participated in the field investigation and identification work, acted as the corresponding author of the paper. Jiang-Hong Yu participated in the field investigation and completed part of the data processing and content writing. Feng Liu Collect plant specimens in the field and take photographs for identification. Zheng-Ren Chen participated in field investigation and compared plant specimens to identify species and guide the writing of the paper.
Tian-Rou Wu https://orcid.org/0009-0001-5223-0567
Jian Xu https://orcid.org/0000-0002-0714-0917
Ming-Tai An https://orcid.org/0000-0003-3886-0287
Jiang-Hong Yu https://orcid.org/0000-0003-1765-8557
All of the data that support the findings of this study are available in the main text.