Research Article |
Corresponding author: Jian-Wen Shao ( shaojw@ahnu.edu.cn ) Academic editor: Lorenzo Peruzzi
© 2022 Si-Yu Zhang, Hao-Tian Wang, Ying-Feng Hu, Wei Zhang, Song Hu, Jian-Wen Shao.
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:
Zhang S-Y, Wang H-T, Hu Y-F, Zhang W, Hu S, Shao J-W (2022) Lycoris insularis (Amaryllidaceae), a new species from eastern China revealed by morphological and molecular evidence. PhytoKeys 206: 153-165. https://doi.org/10.3897/phytokeys.206.90720
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Lycoris insularis S.Y.Zhang & J.W.Shao, a new fertile diploid species from coastal provinces in eastern China is described. This new species is most similar to L. sprengeri in morphology and has been misidentified as the latter for a long time. However, it can be distinguished from the latter by the relatively longer perianth tube (1.5‒2.5 cm vs. less than 1.3 cm), a characteristic that was overlooked before. Phylogenetic analysis, based on complete plastid genome, showed that L. insularis is not genetically related to L. sprengeri in the genus. The former was a sister group of L. sanguinea, while the latter was closely related to L. longituba and L. chinensis and they were respectively located on different clades that were separated at the base of the phylogenetic tree. The chromosome number of L. insularis is 2n = 22. At present, as the new species is relatively widely distributed and the wild population can normally reproduce by seeds, we evaluate it as LC (Least Concern) according to criteria of the IUCN Red List.
Lycoris sprengeri, taxonomy, chloroplast genome, cryptic species
Lycoris Herb. (Amaryllidaceae) is a typical pan-East Asian genus, currently considered to have about 9 fertile diploid species, mainly distributed in China, Japan and South Korea (
Lycoris sprengeri Comes ex Baker was first described in 1902, being only distributed in China. The type specimen was collected from around Xiangyang City, Hubei Province in Central China (
Recently, we collected some populations of L. sprengeri from inland areas, including Guangshui City, Hubei Province (near the collection location of the type specimen) and the surrounding areas of Dabie Mountains in Anhui Province. In the field, we noticed that there was a certain difference in morphology between those plants (identified as L. sprengeri) occurring inland and those from the coastal area: the flower colour of the former varied from white to dark purple in a population and the length of the perianth tube is short (less than 1.3 cm), while the flower colour of the latter is relatively stable, generally pink and the perianth tube is long (usually about 2.0 cm). After further morphological observations, chloroplast sequence alignment and phylogenetic analysis and checking the related type specimens, we recognised that the plants found in the inland area were the genuine L. sprengeri, while those plants collected from the coastal area (identified as L. sprengeri) are actually an undescribed new species. Thus, we named it as Lycoris insularis S.Y.Zhang & J.W.Shao and describe it here.
Bulbs for study and observation were successively collected from the wild in 2016 to 2021. The information for the collection sites is shown in Table
Code | Locations | Altitude | GenBank acc. no |
---|---|---|---|
L. sprengeri (inland area) | |||
01 | Yudian Town, Guangshui City, Suizhou City, Hubei Province | 278 m | OP034616 |
02 | Qiaotou Town, Mingguang City, Chuzhou City, Anhui Province | 91 m | OP034617 |
03 | Yinjian Town, Fengyang County, Chuzhou City, Anhui Province | 137 m | OP034618 |
04 | Huangjia Town, Tongcheng City, Anqing City, Anhui Province | 122 m | OP034619 |
05 | Pingshan Town, Huaining County, Anqing City, Anhui Province | 110 m | OP034620 |
L. insularis (coastal area) | |||
01 | Damao Island, Dinghai District, Zhoushan City, Zhejiang Province | 22 m | OP034614 |
02 | Chunxiao Town, Beilun District, Ningbo City, Zhejiang Province | 8 m | OP034615 |
03 | Shanjuan Town, Yixing City, Wuxi City, Jiangsu Province | 42 m | ON611639 |
L. chinensis | |||
01 | Bozhou Town, Xinhuang County, Huaihua City, Hunan Province | 542 m | OP034613 |
DNA samples were extracted from leaves dried by silica gel according to the modified cetyltrimethylammonium bromide (mCTAB) extraction protocol (
All sequences were aligned by MACSE v.2 (
L. insularis and L. sprengeri are very similar in many morphological characteristics, such as the leaves being slightly twisted when young, the apex being lavender and becoming a narrow and a long band when mature; the perianth is mainly pink, the apex is usually pale blue and the perianth lobes are rarely shrivelled, slightly or almost not rolling back. However, after careful observation, we found there are certain differences between them in the perianth tube length and flower colour. In all L. sprengeri populations observed so far, the perianth tube is short (0.5‒1.3 cm) and the flower colour ranged from nearly white (about 2%) to dark purple (about 2%), although most of the flowers are pale pink. In all populations of L. insularis, the perianth tube is relatively long (usually 1.5‒2.5 cm) and the flower colours are almost all either deep or shallow pink (Figs
Comparison and variation of perianth tube length of Lycoris sprengeri (01‒05, filled with orange) and L. insularis (01‒03, filled with magenta). In the boxplot, the horizontal line shows the median, the bottom and top of the box show the first and third quartiles. Boxplots marked with different letters differ significantly (post hoc test, P < 0.05).
The length of complete chloroplast genome of Lycoris insularis comprised 158,641‒159,121 bp and L. sprengeri comprised 158,509‒159,348 bp (Fig.
Basic characteristics of chloroplast genomes of Lycoris insularis and L. sprengeri.
Characteristic | L. insularis | L. sprengeri |
---|---|---|
Total length (bp) | 158,641‒159,121 | 158,509‒159,348 |
GC% | 37.7%‒37.8% | 37.8%‒37.9% |
LSC length (bp) | 86,488‒86,590 | 86,483‒86,941 |
SSC length (bp) | 18,496‒18,544 | 18,469‒18,501 |
IR length (bp) | 26,806‒27,082 | 26,764‒27,018 |
Total genes | 112 | 112 |
Protein-coding genes | 78 | 78 |
rRNA genes | 4 | 4 |
tRNA genes | 30 | 30 |
The phylogenetic relationship, based on the complete chloroplast genome containing eight species, is shown in Fig.
Phylogeny of the fertile diploid Lycoris species, based on the complete chloroplast genome. Numbers above branches are Maximum Likelihood bootstrap values (BS)/Bayesian posterior probability (BPP). The species name with accession numbers were download from NCBI. * showing those sequences which were misidentified as L. sprengeri in NCBI.
China. Zhejiang, Zhoushan City, Dinghai District, Damao Island, 29°56'55.4"N, 122°3'9.46"E, under the broad-leaved forest near the water on the Island, 22 m a.s.l., 18 August 2019, S.Y. Zhang, ZSY201908001 (holotype: ANUB008515!; isotypes: ANUB008516!, ANUB008517!, CSH0192378!, NPH001410!).
The new species resembles Lycoris sprengeri by young leaves swirling and rising, tepals usually pink with blue apex and tepals length substantially similar, but differs in significantly longer perianth tube (1.5‒2.5 cm vs. 0.5‒1.3 cm).
Perennial herb. Bulb subglobose or ovate, 3‒5 cm in diameter, covered brown epidermis, with fine lines on the epidermis. Leaves linear, often 6‒9, blunt apex, appearing in winter or early spring, 40‒60 cm long, 6‒15 mm wide, pale green, mid-rib slightly sunken, covered with a little white powder. Inflorescence scapose, 40‒60 cm high, green or reddish-brown; two spathe bracts, lanceolate, about 3 cm long, 8‒12 mm wide; 5‒7 flowers per umbels, pedicels 1.5‒3 cm long, diameter about 3 mm; flowers usually pink with blue apex, occasionally white, blue or all pink; perianth lobes oblanceolate, 4.5‒7 cm long, about 0.9‒1.5 cm wide, apex slightly reversed; perianth tubes cream or pink, about 15‒25 mm long. Filaments 5‒7 cm long, pink, slightly longer than tepals; anther yellow, 3‒5 mm long; pistil length 6‒10 cm, pink or with purple apex. Ovary 5 mm in diameter, spherical and green. Capsules three-lobed, green or with light brown when mature; seeds black, spherical, about 7 mm in diameter.
Flowering from late July to late September; fruiting in September to October; leaves growing in winter or early spring (December to February of the following year).
Most of the populations grow on the hillsides or island slopes near the sea (the coastal areas of Shanghai City, Zhejiang Province and Fujian Province) and sporadic populations grow around the inland hills and valleys (Fig.
Hăi bīn shí suàn (海滨石蒜).
Latin insula, island, and -aris, belonging to; the specific epithet alludes to occurrence of the new species in Damao Island.
This species can reproduce asexually by duplication of bulbs (about three times per two years) and also can sexually reproduce through seeds, taking about five years from seed germination to flowering.
Compared with other species in Lycoris, L. insularis has a wider distribution area and many populations grow on inaccessible islands. After our field investigation, a large number of bulbs in Yixing and Huzhou were excavated for greening and landscaping, but there are still a large number of L. insularis populations in the wild in other areas. Thus, we classified its conservation status as LC (Least Concern), according to the IUCN Red List Criteria (
China. Jiangsu Province: Yixing City, Shanjuan Cave, 25 Aug 1960, Wen-zhe Fang 00110645 (PE); Shanghai City: Songjiang District, Tianma Mountain, 8 Sept 1963, Guang-jin Fan 00110641 (PE); Songjiang District, Tianma Mountain, alt. 59 m, 14 Sept 2013, Ting Zhang 1393054 (KUN); Songjiang District, Dongshe Mountain, alt. 21 m, 28 Jul 2015, Bin-jie Ge, Tian Li CSH0098245 (CSH); Zhejiang Province: Putuo District, Taohua Island, alt. 19 m, 18 Sept 2017, Yong-jie Guo, Li Huang, Zheng-yu Zuo & Ting Guo 1446199 (KUN); Jiaojiang County, Jiangshan Island, alt. 3 m, 25 Oct 2016, Yong-jie Guo, Qiao-rong Zhang, Li Huang, Lian-yi Li, Pei Li, De-ming He, Ying-hong Yang 1451350 (KUN); Ruian County, Shuangfengshan Island, alt. 46 m, 11 Oct 2017, Yong-jie Guo, Qiao-rong Zhang, Yun-hua Fang, Xing-xu Sun 1450112 (KUN); Putuo District, Putuo Mountain, alt. 62 m, 24 Aug 2015, Bao-cheng Wu NAS00591943, NAS00591944, NAS00591945 (NAS); Putuo District, Xiaomayi Island, 26 Jul 2011 Qi Tian, Zheng-wei Wang CSH0116798 (CSH); Putuo District, Daqing Mountain, alt.18 m, 21 Sept 2015, Bin-jie Ge, Xu Yuan CSH0101312 (CSH); Daishan County, Yushan Village, 21 Sept 2012, Xi-yang Ye CSH0032460 (CSH).
1 | Leaves appear in autumn (Sept‒Oct) | 2 |
– | Leaves appear in winter or spring (Dec‒Feb) | 5 |
2 | Leaves apex obtuse | 3 |
– | Leaves apex acuminate | 4 |
3 | Flowers red, stamen 6‒8 cm | L. radiata |
– | Flowers rose-red, stamen 3‒3.5 cm | L. wulingensis |
4 | Pedicels 15‒22 mm, perianth tube length 1.2‒1.5 cm | L. aurea |
– | Pedicels 8‒9 mm, perianth tube length 1.5‒2 cm | L. traubii |
5 | Leaves width 0.5‒1.5 cm, apex rose-red when young | 6 |
– | Leaves width 1.5‒3 cm, apex always green | 8 |
6 | Flowers orange | L. sanguinea |
– | Flowers pink, apex usually blue | 7 |
7 | Perianth tube length 1.5‒2.5 cm | L. insularis |
– | Perianth tube length 0.5‒1.3 cm | L. sprengeri |
8 | Flowers actinomorphic, perianth tubes 3‒6 cm | L. longituba |
– | Flowers zygomorphous, perianth tubes 1.5‒2.5 cm | 9 |
9 | Leaves width ca. 1.5 cm, tepals orange, apex usually red | L. tsinlingensis |
– | Leaves width 1.5‒2.5 cm, tepals yellow to light orange | L. chinensis |
Based on our research, especially the phylogenetic tree constructed by the complete chloroplast genome (Fig.
The known karyotypes of fertile diploid species in Lycoris can be roughly divided into two categories: 2n = 12‒16 (such as L. traubii, L. aurea, L. chinensis and L. longituba) or 2n = 22 (such as L. sanguinea, L. radiata, L. wulingensis and L. sprengeri) (
Thanks to Zhang-jie Huang, Cheng-sheng Li and Miao Gao for their help in writing the first draft. Thanks to Yao Zhu and De-ying Liu for their help in the collection of experimental materials. This work was supported by National Natural Science Foundation of China (32070370) and China’s National Basic Science and Technology Program (2019FY101810).