Research Article |
Corresponding author: Lang Li ( lilang@xtbg.ac.cn ) Corresponding author: Jie Li ( jieli@xtbg.ac.cn ) Academic editor: Andrés E. Ortiz-Rodriguez
© 2024 Lang Li, Dian-Yang Zou, Ya-Meng Mao, Jie Li.
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:
Li L, Zou D-y, Mao Y-m, Li J (2024) Alseodaphnopsis maguanensis is conspecific with A. hokouensis (Lauraceae) based on morphological and molecular evidence. PhytoKeys 242: 281-291. https://doi.org/10.3897/phytokeys.242.115679
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Based on both morphological and molecular evidence, it is confirmed that Alseodaphnopsis maguanensis is conspecific with A. hokouensis. Hence, Alseodaphnopsis maguanensis is treated as a synonym of A. hokouensis here. The conservation status of Alseodaphnopsis hokouensis is also re-evaluated according to the IUCN Red List Categories and Criteria in this study.
Alseodaphnopsis, Lauraceae, morphology, phylogeny, synonym, taxonomy
As one of the largest basal angiosperm families, Lauraceae includes more than 50 genera and 2500 ~ 3000 species distributed in tropical and subtropical regions worldwide (
Alseodaphnopsis hokouensis (H. W. Li) H. W. Li & J. Li was first published as an Alseodaphne Nees species (
During recent field surveys in SE Yunnan (SW China), some flowering and fruiting individuals of Alseodaphnopsis hokouensis were found and they resembled a recently-published Alseodaphnopsis species, A. maguanensis L. Li & J. Li (
We conducted field surveys in both Hekou (Hokou) County and Maguan County (Yunnan Province, China) from 2020 to 2022. Flowering specimens of Alseodaphnopsis hokouensis and A. maguanensis were collected from May to June and fruiting specimens were collected from July to September.
Morphological characters of Alseodaphnopsis hokouensis and A. maguanensis were examined and compared in detail, based on fresh and preserved materials as well as dried specimens collected in the field. Herbarium specimens of these two species from the Herbaria HITBC, KUN and PE were also examined.
Based on the work of
Species examined in this study, voucher information, collection localities and GenBank accession numbers for ITS and LEAFY sequences.
Taxon | Voucher | Locality | ITS | LEAFY |
---|---|---|---|---|
Ingroups | ||||
Alseodaphne (4) | ||||
A. gigaphylla Kosterm. | Arifiani DA657 (BO) | Indonesia, Java | HQ697181 | HQ697004 |
A. gracilis Kosterm. | Li L. 20070187 (HITBC) | China, Yunnan | HQ697187 | HQ697036 |
A. huanglianshanensis H. W. Li & Y. M. Shui | Li L. 20080006 (HITBC) | China, Yunnan | HQ6971812 | HQ697007 |
A. semecarpifolia Nees | Arifiani DA658 (BO) | Indonesia, Java | HQ6971814 | HQ697015 |
Alseodaphnopsis (8) | ||||
A. andersonii (King ex Hook. f.) H. W. Li & J. Li, | Li J. & Li L. 20070074 (HITBC) | China, Yunnan | FM957793 | HQ697002 |
A. hainanensis (Merr.) H. W. Li & J. Li | Li L. & Wang Z. H. JFL07 (HITBC) | China, Hainan | MG188587 | MG188634 |
Li L. & Wang Z. H. LMS10 (HITBC) | China, Hainan | MG188586 | MG188633 | |
A. hokouensis (H. W. Li) H. W. Li & J. Li | Li L. et al. 2020064 (HITBC) | China, Yunnan | PP736795 | PP737831 |
Li L. et al. 2020071 (HITBC) | China, Yunnan | PP736796 | PP737832 | |
Li L. et al. 2020072 (HITBC) | China, Yunnan | PP736797 | PP737833 | |
A. maguanensis L. Li & J. Li | Li L. et al. GLQ45 (HITBC) | China, Yunnan | MN906900 | MN906896 |
Li L. et al. GLQ46 (HITBC) | China, Yunnan | MN906901 | MN906897 | |
Li L. et al. 2020086 (HITBC) | China, Yunnan | PP736798 | PP737834 | |
Li L. et al. 2020091 (HITBC) | China, Yunnan | PP7367999 | PP737835 | |
A. petiolaris (Meisn.) H. W. Li & J. Li | Chen J. Q. 07003 (HITBC) | China, Yunnan | FM957796 | HQ697008 |
A. putaoensis L. Li, Y. H. Tan & J. Li | Li L. & Ma H. MM254 (HITBC) | Myanmar, Kachin | MN906902 | MN906898 |
Li L. & Ma H. MM266 (HITBC) | Myanmar, Kachin | MN906903 | MN906899 | |
A. rugosa (Merr. & Chun) H. W. Li & J. Li, | Li L. & Wang Z. H. MYH02 (HITBC) | China, Hainan | MG188585 | MG188635 |
Li L. & Wang Z. H. MYH08 (HITBC) | China, Hainan | MG188584 | MG188640 | |
A. sichourensis (H. W. Li) H. W. Li & J. Li | Song Y. 33225 (HITBC) | China, Yunnan | MG188597 | MG188626 |
A. ximengensis H.W. Li & J. Li | Li J. W. 1235 (HITBC) | China, Yunnan | MG188591 | MG188599 |
Dehaasia (1) | ||||
D. hainanensis Kosterm. | Li L. & Wang Z. H. 20070373 (HITBC) | China, Hainan | FJ719308 | HQ697026 |
Machilus (8) | ||||
M. duthiei King ex Hook. f. | Zhong J. S. 2006094 (HITBC) | China, Yunnan | FJ755425 | HQ697055 |
M. gongshanensis H. W. Li | Chen J. Q. 07002 (HITBC) | China, Yunnan | FJ755416 | HQ697047 |
M. grijsii Hance | Chen J. Q. et al. 2006028 (HITBC) | China, Guangdong | FJ755420 | HQ697049 |
M. kwangtungensis Yang | Chen J. Q. et al. 2006027 (HITBC) | China, Guangdong | FJ755424 | HQ697051 |
M. monticola S. Lee | Li L. & Wang Z. H. 20070323 (HITBC) | China, Hainan | FJ755418 | HQ697057 |
M. platycarpa Chun | Chen J. Q. et al. 2006073 (HITBC) | China, Guangdong | FJ755421 | HQ697067 |
M. robusta W. W. Sm. | Li J. 2002116 (HITBC) | China, Guangxi | FJ755426 | HQ697071 |
M. yunnanensis Lec. | Zhong J. S. 2006093 (HITBC) | China, Yunnan | FJ755415 | HQ697084 |
Nothaphoebe (1) | ||||
N. umbelliflora (Blume) Blume | Arifiani DA495 (BO) | Indonesia, Java | HQ697191 | HQ697088 |
Phoebe (6) | ||||
P. chekiangensis C. B. Shang | Li J. & Li L. 20070188 (HITBC) | China, Zhejiang | FJ755407 | HQ697128 |
P. cuneata (Blume) Blume | Arifiani 40 (MO) | Indonesia, Java | HQ697202 | HQ697130 |
P. formosana (Hayata) Hayata | Rohwer 156 (MJG) | Germany, Bonn | HQ697205 | HQ697136 |
P. lanceolata (Wall. ex Nees) Nees | Chen J. Q. et al. 2006093 (HITBC) | China, Guangdong | FJ755410 | HQ697141 |
P. glaucifolia S. K. Lee & F. N. Wei | Chen J. Q. et al. 2005002 (HITBC) | China, Yunnan | FJ755409 | HQ697150 |
P. neurantha (Hemsl.) Gamble | Li J. & Li L. 20070214 (HITBC) | China, Zhejiang | HQ697209 | HQ697151 |
Outgroups | ||||
Actinodaphne (1) | ||||
A. trichocarpa C. K. Allen | Li L. 20070282 (HITBC) | China, Sichuan | HQ697214 | HQ697166 |
Lindera (1) | ||||
L. erythrocarpa Makino | Li J. & Li L. 20070203 (HITBC) | China, Zhejiang | HQ697215 | HQ697167 |
Litsea (1) | ||||
L. auriculata Chien & Cheng | Li J. & Li L. 20070195 (HITBC) | China, Zhejiang | HQ697217 | HQ697174 |
Neolitsea (1) | ||||
N. howii C. K. Allen | Li L. & Wang Z. H. 20070379 (HITBC) | China, Hainan | HQ697220 | HQ697178 |
The combined dataset including ITS and LEAFY intron II sequences was used for phylogenetic analysis according to the works of
During our field surveys in SE Yunnan, two populations of Alseodaphnopsis hokouensis were found in Huayudong and Qincaitang, Nanxi Town, Hekou County (Fig.
The MP and BI analyses of the ITS + LEAFY intron II combined dataset generated congruent topologies. The Bayesian consensus tree with MP bootstrap (BS) and Bayesian posterior probability (PP) values is shown in Fig.
Bayesian consensus tree of ITS + LEAFY intron II combined dataset. MP bootstrap (BS ≥ 50%) and Bayesian posterior probability (PP ≥ 0.95) values are shown above branches. Act. = Actinodaphne, Al. = Alseodaphne, Als.= Alseodaphnopsis, Deh. = Dehaasia, Lin. = Lindera, Lit. = Litsea, Mac. = Machilus, Neo. = Neolitsea, Not. = Nothaphoebe, Pho. = Phoebe.
The morphological characters of Alseodaphnopsis hokouensis, A. maguanensis and closely related A. rugosa were compared in detail, based on the data from type specimens and protologues (
Comparison of morphological characters, phenologies, habitats and distributions of Alseodaphnopsis hokouensis, A. maguanensis and A. rugosa.
A. hokouensis | A. maguanensis | A. rugosa | |
---|---|---|---|
Morphological characters | |||
Branchlet | terete, brownish when dry, striate, glabrous | terete, greyish, wrinkled, glabrous, with lenticels and leaf scars | terete, robust, wrinkled, with dense leaf scars near top |
Terminal bud | subglobose, bud scales broadly ovate, acute at apex, glabrous | subglobose, bud scales broadly ovate, acute at apex, glabrous | |
Leaf arrangement | leaves alternate | leaves clustered at apex of branchlet, alternate or subverticillate | leaves inserted at apex of branchlet, dense and nearly verticillate |
Petiole | 1.5–3 cm, concave-convex, glabrous | 1.5–2.5 cm, concave-convex, glabrous | robust, 1.5–2.5 cm, glabrous |
Leaf blade | elliptic to oblong, 10.5–17 × 4–6.5 cm, subleathery, glabrous on both surfaces, base broadly cuneate to subrounded, apex abruptly shortly acuminate | oblong-obovate or oblong-oblanceolate, 12–32 × 3.5–9 cm, leathery, glabrous on both surfaces, base cuneate, apex shortly acuminate | oblong-obovate or oblong-oblanceolate, 15–36 × 4–10 cm, leathery, glabrous on both surfaces, base cuneate, apex shortly acuminate. |
Leaf veins | mid-rib elevated abaxially, impressed adaxially, lateral veins 9–13 pairs, transverse veins and veinlets densely reticulate, conspicuous on both surfaces when dry | mid-rib elevated abaxially, impressed adaxially, lateral veins 8–12 pairs, veins and veinlets conspicuous, reticulate, elevated on both surfaces when dry | mid-rib conspicuously elevated abaxially, impressed adaxially, veins and veinlets conspicuous, reticulate |
Inflorescence | panicles subterminal or inserted on lower part of young branchlet, 10.5–15 cm, peduncle and rachis glabrous, pedicels 3–4 mm, glabrous | panicles subterminal, 15–20 cm, peduncle glabrous, pedicels 5–8 mm, glabrous | not seen |
Perianth lobes | perianth lobes 6, ovate, slightly acute, subequal, ca. 2 × 1.5 mm, glabrous outside, grey pubescent inside, deciduous | perianth lobes 6, broadly ovate, acute, subequal, outer ones ca. 2 × 1.5 mm, inner ones ca. 2.5 × 2 mm, glabrous outside, white pubescent inside, deciduous | not seen |
Fertile stamens | fertile stamens 9, minute, ca. 1.5 mm in 1st and 2nd whorls, ca. 1.7 mm in 3rd whorl; filaments villous, ca. 0.7 mm in 1st and 2nd whorls, ca. 1 mm in 3rd whorl, those of 3rd whorl each with 2 stalked orbicular-reniform glands at base, others glandless; anthers of 1st and 2nd whorls oblong, almost as long as filament, with introrse cells, those of 3rd whorl rectangular, with extrorse cells | fertile stamens 9, ca. 2 mm in 1st and 2nd whorls, ca. 2.2 mm in 3rd whorl; filaments villous, those of 3rd whorl each with 2 shortly-stalked orbicular-cordate glands at base, others glandless; anthers of 1st and 2nd whorls ovate, almost as long as filament, cells all introrse, those of 3rd whorl elliptic, with extrorse cells | not seen |
Staminodes | not seen | ca. 1.5 mm, sagittate, stalked | not seen |
Pistil | ovary ovoid, ca. 1.5 mm, glabrous, attenuate into a ca. 0.5 mm long style; stigma discoid, slightly lobed | ovary ovoid, ca. 1.2 mm, glabrous, attenuate into a ca. 0.8 mm long style; stigma discoid, inconspicuous | not seen |
Infructescence | not seen | subterminal, 10–18 cm, robust, glabrous, with only one or two well-developed fruits | subterminal, ca. 12.5 cm, robust, glabrous |
Fruit | not seen | oblate, 4–5 × 5–6 cm, brown when mature; fruit stalk robust, 3–4 mm in diameter, apex dilated, 5–10 mm in diameter, sometimes nearly cylindrical, fleshy and warty when fresh | oblate, ca. 2.5 × 3 cm deep purple or black when mature; fruit stalk robust, 5–8 mm in diameter on top, fleshy, red and warty when fresh |
Phenology | flowering in May; fruit unknown | flowering in May–Jun; fruiting in Jul–Sep | flower unknown; fruiting in Jul–Dec (fruits mostly found in Oct–Dec) |
Habitat | evergreen broad-leaved forests; ca. 700 m alt. | tropical montane forests in valleys; ca. 800 m alt. | mixed forests in valleys; 1200–1300 m alt. |
Distribution | SE Yunnan, China | SE Yunnan, China | Hainan, China |
To sum up, the results of both morphological comparison and phylogenetic analysis showed that Alseodaphnopsis maguanensis is conspecific with A. hokouensis. Therefore, we propose that Alseodaphnopsis maguanensis should be treated as a synonym of A. hokouensis. The insufficient specimen collection of Alseodaphnopsis hokouensis, especially the lack of fruiting specimens, hinders a comprehensive understanding of the species. The same situation is found in many other rare and endemic Lauraceae species in SE Yunnan, such as species of Alseodaphne, Beilschmiedia Nees, Caryodaphnopsis Airy Shaw, Cryptocarya R. Br., Endiandra R. Br., Machilus Rumph. ex Nees, Phoebe Nees and Syndiclis Hook. f., etc. (
Alseodaphne hokouensis H. W. Li, Act Phytotax. Sin. 17 (2): 71. 1979. Basionym. Type: CHINA. Yunnan Province: Hekou County, 10 May 1953, K. H. Tsai 1039 (holotype: KUN [KUN0047456!]; isotype: KUN [KUN0108581!]).
= Alseodaphnopsis maguanensis L. Li & J. Li, PhytoKeys 138: 27–39. 2020. syn. nov. Type: CHINA. Yunnan Province: Maguan County, 14 May 2016, Lang Li et al. GLQ26 (holotype: HITBC!).
Trees evergreen, up to 20 m tall. Branchlets terete, 3–6 mm in diameter, greyish, striate, glabrous, with lenticels and leaf scars. Terminal buds subglobose, ca. 2 mm in diameter; bud scales broadly ovate, acute at apex, glabrous. Leaves clustered at apex of branchlet, alternate or subverticillate; petiole robust, 2–3 mm thick, 1.5–3 cm long, concave-convex; leaf blade green adaxially, glaucous abaxially when young, but green or pale green when mature, oblong-obovate or oblong-oblanceolate, sometimes elliptic to oblong, 10.5–32 × 3.5–9 cm, subleathery to leathery, glabrous on both surfaces, mid-rib elevated abaxially, impressed adaxially, lateral veins 8–13 pairs, slightly elevated on both surfaces, oblique, evanescent and interconnected near leaf margin, transverse veins and veinlets densely reticulate, conspicuous on both surfaces when dry, base cuneate to broadly cuneate, apex shortly acuminate, sometimes abruptly shortly acuminate. Panicles subterminal, clustered at apex of branchlet, 10.5–20 cm, many-flowered; peduncle branched at middle or above, peduncle and rachis glabrous; bracts and bracteoles linear, ca. 1.5 mm, acute, ciliate, caducous. Pedicels slender, 3–8 mm, slightly dilated on top, glabrous. Flowers small, ca. 2.5 mm. Perianth tube short; perianth lobes 6, broadly ovate, slightly acute, glabrous outside, pubescent inside, subequal, outer ones ca. 2 × 1.5 mm, inner ones ca. 2.5 × 2 mm, deciduous. Fertile stamens 9, minute, 1.5–2 mm in 1st and 2nd whorls, 1.7–2.2 mm in 3rd whorl; filaments villous, 0.7–1 mm in 1st and 2nd whorls, 1–1.2 mm in 3rd whorl, those of 3rd whorl each with 2 shortly stalked orbicular-cordate glands at base, others glandless; anthers of 1st and 2nd whorls ovate, almost as long as filaments, with introrse cells, those of 3rd whorl elliptic, slightly shorter than filaments, with extrorse cells. Staminodes conspicuous, ca. 1.5 mm, sagittate, stalked. Ovary ovoid, 1.2–1.5 mm, glabrous, attenuate into a 0.5–0.8 mm long style; stigma discoid, inconspicuous. Infructescence subterminal, 10–18 cm, robust, glabrous, with one or two well-developed fruits. Fruit oblate, 4–5 × 5–6 cm, immature fruit green, brown when mature, fruit stalk robust, 3–4 mm in diameter, apex dilated, 5–10 mm in diameter, sometimes nearly cylindrical, fleshy and warty when fresh.
Flowering from May to June and fruiting from July to September.
Hekou County and Maguan County, Yunnan Province, China. Tropical limestone forests in valleys, usually near streams, at an elevation of 150–850 m.
Currently, Alseodaphnopsis hokouensis is known from Hekou Country and Maguan Country (Yunnan Province, China) with four populations (Fig.
China. Yunnan Province: Hekou County, 17 June 2020, Lang Li et al. 2020064 (HITBC); 18 June 2020, Lang Li et al. 2020071 & 2020072 (HITBC); 25 August 2020, Lang Li and Guan-long Cao, 2020135, 2020136, 2020137 & 2020138 (HITBC). 13 May 2022, Lang Li and Dian-yang Zou, 2022027 (HITBC). Maguan County, 19 June 2020, Lang Li et al. 2020082, 2020083 & 2020086 (HITBC); 20 June 2020, Lang Li et al. 2020091, 2020092, 2020093, 2020094, 2020095 & 2020096 (HITBC); 24 August 2020, Lang Li and Guan-long Cao, 2020127 & 2020128 (HITBC); 2 August 2022, Lang Li et al. 2022034, 2022035 & 2022036 (HITBC).
The authors are grateful to Xi-bing Guo and Jin-guo Zhang for their kind help during the field surveys and to Jens G. Rohwer for his valuable comments on the manuscript.
The authors have declared that no competing interests exist.
No ethical statement was reported.
This work was financially supported by grants from the National Natural Science Foundation of China (No. 31970222), the Yunnan Fundamental Research Projects (No. 202101AT070067, No. 202201AS070055) and the 14th Five-Year Plan of Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences (XTBG-1450101).
Conceptualization: JL, LL. Data curation: LL, DZ. Formal analysis: DZ, LL. Funding acquisition: JL, LL. Investigation: LL, DZ, YM. Methodology: LL. Project administration: JL, LL. Supervision: JL, LL. Visualization: YM, LL. Writing - original draft: LL. Writing - review and editing: JL, DZ, YM, LL.
All of the data that support the findings of this study are available in the main text.