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Research Article
Alseodaphnopsis maguanensis is conspecific with A. hokouensis (Lauraceae) based on morphological and molecular evidence
expand article infoLang Li, Dian-Yang Zou§, Ya-Meng Mao§, Jie Li
‡ Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, Yunnan, China
§ University of Chinese Academy of Sciences, Beijing, China
Open Access

Abstract

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.

Key words

Alseodaphnopsis, Lauraceae, morphology, phylogeny, synonym, taxonomy

Introduction

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 (Chanderbali et al. 2001). The delimitation, delineation and identification of Lauraceae species, especially for tropical species, are always complicated due to limited variation in floral and other morphological characters and insufficient specimen collection (van der Werff and Richter 1996; Rohwer 2000; Li and Li 2004; Yang and Liu 2015).

Alseodaphnopsis hokouensis (H. W. Li) H. W. Li & J. Li was first published as an Alseodaphne Nees species (Lee et al. 1979) and then it was transferred to Alseodaphnopsis H. W. Li & J. Li, a recently described genus of the Lauraceae (Mo et al. 2017). Since it was first collected in 1953, only three herbarium records (including two type specimens) of Alseodaphnopsis hokouensis can be found and no fruiting specimen has been collected.

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 (Li et al. 2020). Based on both morphological and molecular evidence, we confirm that Alseodaphnopsis maguanensis is conspecific with A. hokouensis. Therefore, we treat Alseodaphnopsis maguanensis as a synonym of A. hokouensis here. Based on the field survey data, we also re-evaluate the conservation status of Alseodaphnopsis hokouensis in this study.

Materials and methods

Field surveys

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 studies

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.

Molecular studies and phylogenetic analyses

Based on the work of Li et al. (2020), three individuals of Alseodaphnopsis hokouensis and two additional individuals of A. maguanensis were sampled in the present study. DNA extraction, fragment amplification and sequencing, as well as DNA sequence alignment, followed the work of Li et al. (2020). The newly-obtained DNA sequences were integrated into the ITS + LEAFY intron II combined dataset of the work by Li et al. (2020). Species examined in this study, voucher information, collection locality and GenBank accessions for ITS and LEAFY intron II sequences are given in Table 1.

Table 1.

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 Li et al. (2011), Mo et al. (2017) and Li et al. (2020). Phylogenetic analyses were performed using the Maximum Parsimony (MP) and Bayesian Inference (BI) methods. The MP analysis was performed using PAUP* 4.0b10 (Swofford 2003). The BI analysis was performed using MrBayes v.3.2.6 (Ronquist and Huelsenbeck 2003). Different DNA sequences were defined as separate data partitions. The evolutionary model for each partition (ITS: GTR+I+G; LEAFY intron II: HKY+G) was estimated using jModelTest v.2.1.10 (Darriba et al. 2012) with the Akaike Information Criterion (AIC) (Akaike 1974; Posada and Buckley 2004). The parameters used in both MP and BI analysis followed the work of Li et al. (2020).

Results and discussion

During our field surveys in SE Yunnan, two populations of Alseodaphnopsis hokouensis were found in Huayudong and Qincaitang, Nanxi Town, Hekou County (Fig. 1), each with about 10 mature individuals. No individual was found at the type locality of Alseodaphnopsis hokouensis, Masike, Nanxi Town, Hekou County. Two populations of Alseodaphnopsis maguanensis, each with about 20 mature individuals, were also investigated in Gulinqing Provincial Natural Reserve (Maguan County) according to the work of Li et al. (2020).

Figure 1. 

The recorded localities of Alseodaphnopsis hokouensis and A. maguanensis in SE Yunnan, SW China.

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. 2. As in the work of Li et al. (2020), all Alseodaphnopsis specimens sampled in the present study formed a well-defined clade (BS 88%, PP 1.00). Within the Alseodaphnopsis clade, there are two well-supported subclades which consist of four and five species, respectively. All individuals of Alseodaphnopsis hokouensis and A. maguanensis formed a strongly-supported clade (BS 100%, PP 1.00), closely related to A. rugosa. Within this clade, the individuals of these two species are mixed with each other and their relationships are poorly resolved.

Figure 2. 

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 (Li et al. 2008; Li et al. 2020). Morphological comparisons showed that Alseodaphnopsis maguanensis is almost the same as A. hokouensis, except that the leaves of the former are oblong-obovate or oblong-oblanceolate and those of the latter are elliptic to oblong (Table 2). After carefully checking the specimens collected from field surveys, we found that the leaves of both species can vary from elliptic, oblong to oblong-obovate or oblong-oblanceolate (Fig. 3). They are usually oblong-obovate or oblong-oblanceolate and occasionally elliptic to oblong. According to the newly-obtained fruiting specimens of Alseodaphnopsis hokouensis, the fruit characters of Alseodaphnopsis hokouensis are also consistent with those of A. maguanensis (Fig. 3). Additionally, the phenologies and habitats of these two species are quite similar and their distribution areas are adjacent as well (Fig. 1).

Table 2.

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
Figure 3. 

Alseodaphnopsis hokouensis A flowering branchlet B fruiting branchlet C different shapes of leaf blades. Photographed by Lang Li.

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. (Li et al. 2008). Thus, more field surveys and specimen collections are suggested in SE Yunnan in order to improve the understanding of these rare and endemic Lauraceae species.

Taxonomic treatment

Alseodaphnopsis hokouensis (H. W. Li) H. W. Li & J. Li

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!).

Revised description

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.

Phenology

Flowering from May to June and fruiting from July to September.

Distribution and habitat

Hekou County and Maguan County, Yunnan Province, China. Tropical limestone forests in valleys, usually near streams, at an elevation of 150–850 m.

Conservation status

Currently, Alseodaphnopsis hokouensis is known from Hekou Country and Maguan Country (Yunnan Province, China) with four populations (Fig. 1). Two populations found in Maguan Country are all located in Gulinqing Provincial Nature Reserve, each with about 20 mature individuals (Fig. 4). The other two populations found in Hekou Country, each with about 10 mature individuals, are not located in any nature reserve and those individuals mostly occur on the roadsides or in strongly-disturbed forests near the villages (Fig. 4). No individual was found at the type locality, Masike, Nanxi Town, Hekou County. According to the IUCN Red List Categories and Criteria version 15.1 (July 2022), the conservation status of Alseodaphnopsis hokouensis is re-evaluated as Critically Endangered (CR, C2a(i)).

Figure 4. 

Different habitats of Alseodaphnopsis hokouensis A in the forest of the Nature Reserve B on the roadside C in the strongly disturbed forest near the village.

Additional specimens examined

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).

Acknowledgements

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.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

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).

Author contributions

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.

Author ORCIDs

Lang Li https://orcid.org/0000-0002-7439-2030

Jie Li https://orcid.org/0000-0001-8067-749X

Data availability

All of the data that support the findings of this study are available in the main text.

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