Login
Register
Submit a Paper
Propose a Special lssue
Open Access
ARTICLE
Taxonomic Clarification and Revision of Lycoris haywardii Traub
1 Hangzhou Botanical Garden, Hangzhou West Lake Academy of Landscape Science, Hangzhou, China
2 College of Life Sciences, Anhui Normal University, Wuhu, China
3 College of Civil and Architecture Engineering, Chuzhou University, Chuzhou, China
4 Institute of Botany, Jiangsu Province and Chinese Academy of Sciences, Nanjing, China
5 Jiangsu Key Laboratory for Conservation and Utilization of Plant Resources, Nanjing, China
* Corresponding Author: Yuhong Zheng. Email:
(This article belongs to the Special Issue: Taxonomy, Phytogeography and Ecology of Mediterranean Flora)
Phyton-International Journal of Experimental Botany 2026, 95(4), 20 https://doi.org/10.32604/phyton.2026.075166
Received 26 October 2025; Accepted 21 April 2026; Issue published 28 April 2026
View Full Text
Download PDFAbstract
Based on an integrative analysis of morphological traits, phenological characteristics, karyotype, and chloroplast genome data, we revised the taxonomic status of Lycoris haywardii Traub, reclassifying it as a cultivar of Lycoris rosea Traub ex Moldenke, designated as Lycoris rosea ‘Lin’. Lycoris haywardii Traub was originally published by Traub in 1957 as a spring-leafing taxon with orchid purple flowers and stamens slightly shorter than the perianth, with a flowering period from July to mid-August and a purported distribution in Japan. However, populations cultivated in Hangzhou Botanical Garden, previously referred to as L. haywardii, exhibited autumnal leaf emergence, deep purple-pink flowers, and stamens longer than the perianth. Previous artificial hybridization experiments and our current chloroplast genome analyses, along with detailed morphological and phenological observations, confirmed that it is not conspecific with the original L. haywardii. In addition, they are natural hybrids derived from L. radiata var. pumila Grey and L. insularis S. Y. Zhang & J. W. Shao, the same as the origin of L. rosea. Therefore, we formally revised the taxonomy and provided a detailed morphological description of this cultivar.Keywords
The genus Lycoris Herb., belonging to the family Amaryllidaceae, is primarily distributed across East Asia, including China, Japan, and the Korean Peninsula. Scattered populations also occur in parts of Southeast Asia (Laos, Myanmar, Thailand, and Vietnam) and South Asia (India and Pakistan) [1]. To date, a total of 31 species and 5 varieties of Lycoris have been validly published worldwide. The highest species diversity is in eastern China, which is regarded as the center of diversity for the genus [2,3]. Most Lycoris species were described prior to the 1950s by Western botanists based on cultivated or imported specimens, without comprehensive or systematic investigation of wild populations. Many species lacked the designation of type specimens at the time of publication (before 1958); and where types were designated, most are now untraceable.
Lycoris haywardii Traub was published by Traub in 1957. It was characterized by spring-emerging leaves, floral coloration darker than that of L. sprengeri Comes ex Baker but lighter than that of L. aurea, and stamens slightly shorter than the perianth. Flowering occurs from July to mid-August, approximately two weeks earlier than L. sprengeri. Its original distribution was reported as Japan [4]. The designated specimens include No. 554 (holotype, TRA), collected by Sam Caldwell in Nashville, Tennessee, and No. 291 (paratype, TRA), collected by Wyndham Hayward in Winter Park, Florida.
Hsu et al. [5] revised the taxonomic identity of a cultivated population maintained at the Hangzhou Botanical Garden, reporting an autumnal leaf emergence rather than spring. The material studied was of cultivated origin only. In addition, Hsu et al. [5] observed that the stamens were clearly exserted beyond the perianth, traits not consistent with Traub’s original description. According to Adams [6], the distribution of L. haywardii was reported in both China and Japan. However, the species is not included in either Flora Reipublicae Popularis Sinicae [7] or Flora of China [1]. Within the taxonomy of Lycoris, leaf emergence phenology and floral morphology are considered key diagnostic characters. The traits described by Hsu et al. [5] in the Hangzhou cultivated population differ significantly from those originally documented by Traub [4].
In this study, we conducted morphological observations, karyotype analysis, and chloroplast genome sequencing to clarify the taxonomic identity of the population described by Hsu et al. [5] cultivated in Hangzhou Botanical Garden, providing a scientific basis for the rational conservation and utilization of this genetic resource. Based on our findings, we propose that this population represents a cultivar of L. rosea and designate it as L. rosea ‘Lin’.
Bulbs of L. rosea ‘Lin’ were collected in Hangzhou Botanical Garden which is cultivated abundantly nowadays. Hangzhou Botanical Garden is located at 30°15′34″ N, 120°07′15″ E, a typical subtropical monsoon climate, and an average altitude is 20 m. Other materials used for karyotype analysis or molecular phylogenetic studies were collected from a wild population located in Chunxiao Town, Ningbo City, Zhejiang Province (29°47′33″ N, 121°57′07″ E), and introduced for cultivation in Hangzhou Botanical Garden.
2.2.1 Morphology and Chromosome Observations
Morphological traits examined in this study included: bulb size, presence of a pale central band on the leaf blade, leaf emergence phenology, leaf color, leaf length, leaf width (the widest site), leaf apex shape, flower morphology, flower color, flowering period, scape height, spathe length, number of individual flowers per umbel, perianth length, perianth width, perianth tube length, stamen length, pistil length, flower diameter, and fruit set. For each trait, measurements were taken from ten individual plants per taxon.
2.2.2 Acquisition, Annotation, Comparison and Phylogenetic Analysis of Chloroplast Genome
Genome DNA was extracted from fresh leaves of two individuals of L. rosea ‘Lin’ (Voucher No. HZ077536, HHBG) and one individual of wild L. rosea using the modified CTAB (mCTAB) method [8,9]. After Polymerase Chain Reaction (PCR), we used a NanoDrop 1000 Spectrophotometer and agarose gel electrophoresis to check the DNA quality. DNA library building, Genome Skimming and FastQC were outsourced to Wuhan Frasergen Genetic Information Co., Ltd. (Wuhan, China), which employed Illumina HiSeq 6000 for analysis. In this study, the 3 Gb raw data generated by genome skimming were assembled and analyzed. With Lycoris insularis (OP034614) as the reference, the complete chloroplast genome sequences were assembled using GetOrganelle v.1.7.1, followed by genome annotation performed with PGA [10,11,12]. Finally, the annotated sequences were submitted to the National Center for Biotechnology Information (NCBI) with the accession numbers PV839568–PV839570 (Fig. 1). This study was performed on the eleven complete chloroplast genomes in Lycoris Herb., of which ten were downloaded from NCBI. Narcissus poeticus (MH706763) was selected as the outgroup. All sequences were aligned by MACSE v.2 [13]. Maximum Likelihood (ML) and Bayesian Inference (BI) methods were used to determine the phylogenetic relationships, which the best-fit model of DNA substitution being estimated by ModelFinder [14,15]. ML analysis was conducted using the GTR+G+I model with 1000 bootstrap replicates by IQtree v.1.6.8 [16]. Bayesian analysis was constructed with eight independent chains for 1,000,000 generations and sampling every 1000 generations by MrBayes v.3.2.6 [17,18]. All phylogenetic analyses were performed in Phylosuite [18].
Figure 1: Phylogeny of 11 taxa in Lycoris Herb., based on the complete chloroplast genome. Numbers above branches are Maximum Likelihood bootstrap values (BS)/Bayesian posterior probability (BPP), unlabeled branches indicate support values of 100/1. The species name with accession numbers were download from NCBI. *showing those sequences which were misidentified as L. insularis in NCBI [10].
3.1 The Origin of the Lycoris haywardii Bulbs
In the protologue of Lycoris haywardii [4] listed Japan as the place of origin. However, at the beginning of the same publication, Traub noted that Hayward had imported Lycoris bulbs from China between the early 1940s and 1948. After 1945, Hayward and others also imported numerous bulbs from Japan. Although L. haywardii was included among these imported materials, its precise geographic origin was not explicitly stated.
Caldwell [19,20] documented that in October 1948, he received three bulbs labeled “L. incarnata” from Hayward. These bulbs were part of a lot of approximately 100 bulbs purchased from Shanghai, China. After five years of cultivation, the plants flowered. Based on his observations, Caldwell considered this taxon closely related to L. sprengeri, with distinguishing features including less overlap at the base of the perianth segments, reduced recurving at the apex, lighter flower color, and a non-overlapping flowering period. It was from this population that Traub, honoring Hayward, formally described L. haywardii.
Hsu et al. [5] revised the taxonomic identity of L. haywardii based on specimens cultivated in Hangzhou Botanical Garden (voucher specimens: J. Z. Lin 002, 003; Z. Z. Yu 026). In agreement with Traub’s assertion, they recognized the taxon as of cultivated origin and revised its distribution to “cultivated only.” However, the voucher specimens cited in Hsu et al.’s revision are currently untraceable, and the evidence supporting the amended distribution remains unverified.
Lin et al. [21] reported results from an artificial hybridization experiment conducted in 1984 between L. radiata var. pumila and L. sprengeri. The F1 generation exhibited phenotypic traits highly similar to L. rosea. Morphological, phenological, vegetative (leaf shape, leaf color, bulb morphology), and karyotypic characteristics of L. rosea ‘Lin’ were consistent with those of L. rosea [22]. This supports the hypothesis that both taxa likely originated from a natural hybridization event between L. radiata var. pumila and L. sprengeri. This may have been a key reason why Hsu et al. [5] revised the distribution of L. haywardii Traub to cultivated status only.
Notably, according to Lin et al. [23] and Lin [24], both L. rosea ‘Lin’ and L. rosea were originally collected from Hangzhou, Zhejiang Province, China.
Table 1 summarizes the comparative morphological characteristics described by Traub [4], Caldwell [19], Hsu et al. [5], and those observed in L. rosea ‘Lin’. Fig. 2 shows photographs taken by Caldwell in 1957 of L. sprengeri and L. haywardii cultivated in Nashville, Tennessee.
Hsu et al. [5] conducted a taxonomic revision of L. haywardii based on a cultivated population at the Hangzhou Botanical Garden. They revised the leaf emergence period from spring (as originally described by Traub) to autumn. Additionally, Hsu et al. noted that the stamens of voucher specimens were prominently exserted beyond the perianth, indicating bilateral floral symmetry. According to Traub [4], L. haywardii exhibits spring-emerging leaves and actinomorphic (radially symmetrical) flowers, while Hsu et al. [5] reported autumn-emerging leaves and zygomorphic (bilaterally symmetrical) flowers in the revised concept of L. haywardii. These differences are taxonomically significant.
Currently, the cultivated L. rosea ‘Lin’ at Hangzhou Botanical Garden (referenced in Table 1) displays a smaller flowering bulb diameter (~2 cm), shorter but broader leaves (30–40 cm in length, 1.2–1.4 cm in width), and a typical bilaterally symmetrical floral morphology. The flower color is darker (deep purple-pink), the scape is taller (60–70 cm), the spathe is longer (4–4.5 cm), and the number of flowers per umbel is greater (6–8). The perianth segments are also longer, measuring approximately 5.5–6 cm.
Table 1: Comparative Morphological Characteristics of L. haywardii and L. rosea ‘Lin’.
| Character | L. haywardii Traub [4,19] | L. haywardii Traub [5] | L. rosea ‘Lin’ |
|---|---|---|---|
| Bulb size | 3/4—1 1/8 inch (1.89–2.86 cm) | — | 2 cm |
| Pale midrib on leaf blade | Absent | — | Absent |
| Leaf emergence period | Late January to early February | Autumn | November |
| Leaf color | Deep green, slightly glaucescent | Deep green, slightly glaucescent | Greyish olive green |
| Leaf length | 46–48 cm | 48 cm | 30–40 cm |
| Leaf width | 8–9 mm (middle part) | 7–11 mm | 12–14 mm |
| Leaf apex shape | Rounded obtuse | — | Rounded obtuse |
| Flower symmetry | Actinomorphic (radially symmetrical) | Actinomorphic | Zygomorphic |
| Flower color | Orchid purple | Reddish violet | Deep purple pink |
| Flowering period | July to mid-August | July–August | July–August |
| Scape height | 46–48 cm | — | 60–70 cm |
| Spathe length | 3.7 cm | — | 4–4.5 cm |
| Number of flowers per umbel | 4–5 | — | 6–8 |
| Perianth length | 4.2–5.5 cm | 4.4–5.5 cm | 5.5–6 cm |
| Perianth width | 1–1.1 cm | 1–1.1 cm | 1.1–1.2 cm |
| Corona tube length | 1.1 cm | 1.1–1.3 cm | 1–1.2 cm |
| Stamen length | Slightly shorter than perianth segments | Slightly shorter than perianth segments | Ca. 1/6 longer than perianth |
| Pistil length | Slightly longer than perianth segments | Slightly longer than perianth segments | Longer than perianth |
| Flower diameter | 2 3/4 inch (7 cm) | — | 6–7 cm |
| Fertility | Fertile | — | Fertile |
Figure 2: L. sprengeri (left) and L. haywardii (right) cultivated by Caldwell in Nashville, Tennessee (photographed in 1957).
3.3 Characteristics of the Complete Chloroplast Genome
The complete chloroplast genome of L. rosea ‘Lin’ was determined to be 158,472–158,478 base pairs in length (Fig. 1 and Fig. 3), with an overall GC content of 37.80%. The genome exhibited a typical quadripartite structure consisting of a pair of inverted repeats (IRs), a large single-copy (LSC) region, and a small single-copy (SSC) region, measuring 26,828 bp, 86,278 bp, and 18,541 bp, respectively. A total of 112 unique genes were annotated, including 78 protein-coding genes, 30 tRNA genes, and 4 rRNA genes.
Using Narcissus poeticus as the outgroup, a phylogenetic tree was reconstructed based on the complete chloroplast genome sequences of L. rosea ‘Lin’ and 11 other Lycoris taxa (Fig. 1). The genus Lycoris was divided into two major clades. The first clade comprised L. aurea, L. longifolia, L. radiata, L. sprengeri, L. chinensis and L. longituba, while the second clade included L. sanguinea, L. rosea, L. rosea ‘Lin’, and L. insularis. Notably, the publicly available L. rosea plastome (GenBank accession: OL960459) clustered with L. rosea, L. rosea ‘Lin’, and L. insularis in a strongly supported monophyletic group (Bootstrap support = 1.00, Bayesian posterior probability = 100).
Given that chloroplast genomes in Lycoris (as in most angiosperms) are maternally inherited [10], the phylogenetic topology suggests that L. insularis is the maternal parent of both L. rosea and L. rosea ‘Lin’.
Figure 3: Plastid genome map of L. rosea ‘Lin’. Genes marked with an asterisk (*) harbor a single intron.
3.4 Taxonomic Revision of Lycoris haywardii
L. rosea was first published in 1949 by Traub and Moldenke [25] in Amaryllidaceae: Tribe Amarylleae, and was classified under subgenus Eulycoris. The authors established this taxon based on the description of a variant group of L. radiata provided by Worsley [26]. The original diagnosis was relatively brief: the leaves are much broader than those of L. radiata, the umbel contains five flowers, the flowers are larger and pink in color, and the perianth segments are ascending and spreading. Chromosome number was unknown, and the distribution was reported as East Asia. No type specimen was designated.
Worsley [26] considered the taxon to possess intermediate traits between L. albiflora and L. straminea, and suggested a probable hybrid origin. Both the Flora Reipublicae Popularis Sinicae and Flora of China have recognized this species, listing its native range as Jiangsu and Zhejiang provinces, with the type locality reported as Shanghai.
Shi et al. [27] conducted sequence comparisons of the internal transcribed spacer (ITS) region of the nuclear genome and provided molecular evidence supporting the hypothesis that both L. rosea and L. rosea ‘Lin’ are hybrid derivatives of L. radiata var. pumila and L. sprengeri.
Comparative analysis of chloroplast genome structures further supports their close relationship. L. rosea ‘Lin’ and L. rosea exhibit identical lengths in their inverted repeat (IR) and small single-copy (SSC) regions, measuring 26,828 bp and 18,541 bp, respectively. They also share an identical GC content of 37.8%. However, significant differences exist in the total number of genes, protein-coding genes, and tRNA gene content. L. rosea harbors 133 genes, including 87 protein-coding genes, 38 tRNA genes, and 4 rRNA genes, whereas L. rosea ‘Lin’ contains 112 genes, with 78 protein-coding genes, 30 tRNA genes, and 4 rRNA genes [28].
Morphologically, L. rosea and L. rosea ‘Lin’ are also remarkably similar, as evidenced by Table 1 and Fig. 4.
Figure 4: L. rosea ‘Lin’ (left) and L. rosea (right).
When Lycoris haywardii was described by Traub in 1957, its habitat was recorded as Japan. However, in the same journal issue, Caldwell provided a detailed morphological and phenological description of the species, stating its origin as Shanghai. Both authors had received their bulbs from the same source, Hayward. Adams [6] later considered L. haywardii to be distributed in both Japan and China. All three sources—Traub [4], Caldwell [19], and Adams [6]—consistently documented spring leaf emergence and stamens that were slightly shorter than the perianth segments.
In contrast, Hsu et al. [5] revised the taxonomic identity of L. haywardii based on a cultivated population at Hangzhou Botanical Garden, which is now known as L. rosea ‘Lin’. They modified the phenology, reporting autumn leaf emergence, restricted the distribution to cultivated material only, and noted prominently exserted stamens extending beyond the perianth, indicating clear morphological divergence. Evidently, the plant materials used by Hsu et al. [5], i.e., L. rosea ‘Lin’, are not conspecific with the taxon described by Traub [4], Caldwell [19], or Adams [6].
The confusion between L. haywardii and L. rosea ‘Lin’ may partly stem from their shared chromosome number and fertility. Bose [29] reported the karyotype of L. haywardii as 2n = 22 = 16B + 2B+ 4C (=22A), including four satellited chromosomes with proximal centromeres. Similarly, Xu et al. [30] and Liu & Xu [31] documented the karyotype of L. rosea ‘Lin’ as 2n = 22, consistent with Bose’s findings. Both taxa are self-fertile. Phylogenetic analyses based on inter-simple sequence repeat (ISSR) molecular markers conducted by Shi et al. [27] also confirmed a close genetic relationship between L. rosea and L. rosea ‘Lin’.
Taken together, morphological, cytological, and molecular evidence all support the conclusion that L. haywardii and L. rosea ‘Lin’ are distinct taxa. Both L. rosea ‘Lin’ and L. rosea are of hybrid origin, likely derived from hybridization between L. radiata var. pumila and L. insularis. Therefore, it is taxonomically more accurate and scientifically justified to regard the populations currently referred to as L. haywardii by Hsu et al. [5] as a cultivar of L. rosea, under the name L. rosea ‘Lin’.
L. rosea ‘Lin’ P. C. Zhang, S. Y. Zhang & Y. H. Zheng, stat. nov.
Herbs perennial. Bulbs subglobe, Ca. 3–4 cm in diam. Leaves appearing in late Autumn, ligulate, Ca. 30 × 0.7–1.3 cm, apex abtuse. Peduncle solid Ca. 60 cm, umbel 4–8 flowered. Spathe-valves 2, lanceolate, Ca. 3.5 × 1.5 cm. Perianth zygomorphic, red-purple and blue at the tip, tube 1–1.3 cm, lobes oblanceolate, Ca. 4.5–5.5 × 1 cm, margin slightly undulate, apex moderately recurved. Stamens distinctly longer than perianth, Ca. 6 cm, filament red. Style long, filiform, stigma capitate, minute. Flowering late July to mid-August, capsule membranous, loculicidally 3-valved after one and a half months of flowering, seeds few, turgid, testa black, opaque.
Distribution: Cultivated in Hangzhou Botanical Garden.
Specimens: Holotype: HHBG077679, Isotype: HHBG077680.
Notes: The epithet of cultivar is in honor of Ms. Lin Jinzhen.
Combined with morphological traits, phenological characteristics, karyotype, and chloroplast genome data, L. haywardii Traub, which is known only from cultivation in Hangzhou Botanical Garden, is revised as L. rosea ‘Lin’. It has red-purple flowers with blue in the tips. As with L. rosea, it is a fall-leaf species with a flowering period from July to mid-August. Artificial hybridization confirmed that it originated from L. radiata var. pumila and L. insularis. Its chromosome number is 2n = 22. Chloroplast genome sequencing showed L. rosea ‘Lin’ and L. rosea exhibit identical lengths in their inverted repeat (IR) and small single-copy (SSC) regions and share an identical GC content.
Acknowledgement:
Funding Statement: This work was supported by grants from the Science and Technology Development Plan Project of Hangzhou West Lake Scenic Area (2025-008).
Author Contributions: Yuhong Zheng contributed to the conception the study and wrote the manuscript; Pengchong Zhang and Siyu Zhang designed and performed the experiments; Yingfeng Hu and Yaying Mo carried out the statistical analysis. All authors contributed to the manuscript’s revision, read, and approved the submitted version. All authors reviewed and approved the final version of the manuscript.
Availability of Data and Materials: The authors confirm that the data supporting the findings of this study are available within the article.
Ethics Approval: Not Applicable.
Conflicts of Interest: The authors declare no conflicts of interest.
References
1. Ji ZH , Meerow AW . Lycoris Herb. In: Wu Z , Raven PH , editors. Flora of China. Beijing, Chian: Science Press; 2000. p. 264– 73. [Google Scholar]
2. Zhang P , Zheng Y , Zhang S , Pang C , Tian L . Lycoris of Amaryllidaceae in China. In: Ma J , editor. China: mother of gardens in the twenty-first century. Beijing, China: Science Press; 2024. p. 58– 97. (In Chinese). [Google Scholar]
3. Wang R . The ecogeography concerning genus Lycoris. J Southwest For Coll. 1990; 1: 41– 8. (In Chinese). [Google Scholar]
4. Traub H . Lycoris haywardii, L. houdyshelii and L. caldwellii. Plant Life. 1957; 24: 42– 28. [Google Scholar]
5. Hsu PS , Kurita S , Yu ZZ , Lin JZ . Synopsis of the genus Lycoris. Sida. 1994; 16( 2): 301– 31. [Google Scholar]
6. Adams P . Lycoris—jewels among the Amaryllids, a gardener’s perspective. Herbertia. 2011; 65: 108– 233. [Google Scholar]
7. Hsu Y , Hu Z , Huang X , Fan Q . Lycoris Herb. In: Delectis Florae Reipublicae Popularis Sinicae Agendae Academiae Sinicae, editor. Flora Reipublicae Popularis Sinicae. Beijing, China: Science Press; 1985. p. 16– 27. (In Chinese). [Google Scholar]
8. Pahlich E , Gerlitz C . A rapid DNA isolation procedure for small quantities of fresh leaf tissue. Phytochemistry. 1980; 19: 11– 3. doi:10.1016/0031-9422(80)85004-7. [Google Scholar] [CrossRef]
9. Larridon I , Walter HE , Guerrero PC , Duarte M , Cisternas MA , Hernández CP , et al. An integrative approach to understanding the evolution and diversity of Copiapoa (Cactaceae), a threatened endemic Chilean genus from the Atacama Desert. Am J Bot. 2015; 102( 9): 1506– 20. doi:10.3732/ajb.1500168. [Google Scholar] [CrossRef]
10. Zhang SY , Wang HT , Hu YF , Zhang W , Hu S , Shao JW . Lycoris insularis (Amaryllidaceae), a new species from Eastern China revealed by morphological and molecular evidence. PhytoKeys. 2022; 206: 153– 65. doi:10.3897/phytokeys.206.90720. [Google Scholar] [CrossRef]
11. Qu XJ , Moore MJ , Li DZ , Yi TS . PGA: a software package for rapid, accurate, and flexible batch annotation of plastomes. Plant Methods. 2019; 15: 50. doi:10.1186/s13007-019-0435-7. [Google Scholar] [CrossRef]
12. Jin JJ , Yu WB , Yang JB , Song Y , DePamphilis CW , Yi TS , et al. GetOrganelle: a fast and versatile toolkit for accurate de novo assembly of organelle genomes. Genome Biol. 2020; 21( 1): 241. doi:10.1186/s13059-020-02154-5. [Google Scholar] [CrossRef]
13. Ranwez V , Douzery EJP , Cambon C , Chantret N , Delsuc F . MACSE v2: toolkit for the alignment of coding sequences accounting for frameshifts and stop codons. Mol Biol Evol. 2018; 35( 10): 2582– 4. doi:10.1093/molbev/msy159. [Google Scholar] [CrossRef]
14. Minh BQ , Nguyen MAT , von Haeseler A . Ultrafast approximation for phylogenetic bootstrap. Mol Biol Evol. 2013; 30( 5): 1188– 95. doi:10.1093/molbev/mst024. [Google Scholar] [CrossRef]
15. Kalyaanamoorthy S , Minh BQ , Wong TKF , von Haeseler A , Jermiin LS . ModelFinder: fast model selection for accurate phylogenetic estimates. Nat Methods. 2017; 14( 6): 587– 9. doi:10.1038/nmeth.4285. [Google Scholar] [CrossRef]
16. Nguyen LT , Schmidt HA , von Haeseler A , Minh BQ . IQ-TREE: a fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Mol Biol Evol. 2015; 32( 1): 268– 74. doi:10.1093/molbev/msu300. [Google Scholar] [CrossRef]
17. Ronquist F , Teslenko M , van der Mark P , Ayres DL , Darling A , Höhna S , et al. MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Syst Biol. 2012; 61( 3): 539– 42. doi:10.1093/sysbio/sys029. [Google Scholar] [CrossRef]
18. Zhang D , Gao F , Jakovlić I , Zou H , Zhang J , Li WX , et al. PhyloSuite: an integrated and scalable desktop platform for streamlined molecular sequence data management and evolutionary phylogenetics studies. Mol Ecol Resour. 2020; 20( 1): 348– 55. doi:10.1111/1755-0998.13096. [Google Scholar] [CrossRef]
19. Caldwell S . Four new Lycoris for American gardens. Plant Life. 1957; 24: 48– 58. [Google Scholar]
20. Caldwell S . Lycorises: a progress report. Am Hortic Mag. 1962; 41( 2): 61– 94. [Google Scholar]
21. Lin JZ , Yu ZZ , Ruan B . Experimental taxonomy of Lycoris Herb. J Hangzhou Bot Gard. 1992; 2: 5– 11. (In Chinese). [Google Scholar]
22. Xu BS , Huang SF . Karyotype analysis of Lycoris rosea Traub ex Moldenke. Acta Phytotaxon Sin. 1984; 22( 1): 46– 8. [Google Scholar]
23. Lin JZ , Yu ZZ , Ruan B . Exploitation and utilization of Lycoris Herb. J Hangzhou Bot Gard. 1988; 1: 21– 6. (In Chinese). [Google Scholar]
24. Lin JZ . Summary of research on introduction, cultivation, and new cultivar breeding of bulbous root flowers. J Hangzhou Bot Gard. 1991; 2: 5– 12. (In Chinese). [Google Scholar]
25. Traub H , Moldenke H . Amaryllidaceae: tribe amarylleae. Standard, CA, USA: American Plant Life Society; 1949. [Google Scholar]
26. Worsley YA . Lycoris: a garden review. Gard Chron. 1928; 84: 169. [Google Scholar]
27. Shi S , Qiu Y , Wu L , Fu C . Interspecific relationships of Lycoris (Amaryllidaceae) inferred from inter-simple sequence repeat data. Sci Hortic. 2006; 110( 3): 285– 91. doi:10.1016/j.scienta.2006.07.011. [Google Scholar] [CrossRef]
28. Fu WF , Xiao T , Zhang YH . Chloroplast genome characteristics and phylogeny of the genus Lycoris (Amaryllidaceae). Subtrop Plant Sci. 2023; 52( 4): 271– 86. (In Chinese). [Google Scholar]
29. Bose S . Cytological investigations in Lycoris I. The somatic chromosomes of L. caldwellii, L. haywardii and L. houdyshelii. Plant Life. 1957; 13: 34– 9. [Google Scholar]
30. Xu B , Huang S , Lin J , Yu Z , Mao Z . Karyotype analysis of Lycoris haywardii Traub and L. sprengeri Comex ex Baker. Collect Res Pap Nanjing Sun Yat-Sen Bot Gard. 1981; 15–19. (In Chinese). [Google Scholar]
31. Liu Y , Xu B . Karyotype study of Lycoris Herb. Acta Phytotaxon Sin. 1989; 27( 4): 257– 64. (In Chinese). [Google Scholar]
Cite This Article
Copyright © 2026 The Author(s). Published by Tech Science Press.This work is licensed under a Creative Commons Attribution 4.0 International License , which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.
Downloads
Citation Tools
