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Two new species of Alseodaphnopsis (Lauraceae) from southwestern China and northern Myanmar: evidence from morphological and molecular analyses
expand article infoLang Li§, Yun-Hong Tan|§, Hong-Hu Meng§, Hui Ma, Jie Li#§
‡ Core Botanical Gardens, Chinese Academy of Sciences, Mengla, China
§ Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, Mengla, China
| Southeast Asia Biodiversity Research Institute, Chinese Academy of Sciences, Nay Pyi Taw, Myanmar
¶ Xishuangbanna Tropical Botanical Garden, Mengla, China
# Core Botanical Gardens, Chinese Academy of Sciences, Mengla , China
Open Access

Abstract

Alseodaphnopsis maguanensis and A. putaoensis, two new species of Alseodaphnopsis (Lauraceae) from southwestern China (Yunnan Province) and northern Myanmar (Kachin State), are here described and illustrated based on both morphological and molecular evidence. They are morphologically similar to Alseodaphnopsis rugosa and phylogenetically closely related to A. rugosa and A. hainanensis respectively. Their preliminary conservation status is also estimated according to the IUCN Red List Categories and Criteria.

Keywords

Kachin state, phylogenetic analysis, taxonomy, tropical montane forest, Yunnan province

Introduction

Alseodaphnopsis H. W. Li & J. Li, including nine species at present, is a recently described new genus of the Lauraceae (Mo et al. 2017a). Mo et al. (2017a) separated Alseodaphnopsis from the traditionally recognized tropical Asian genus Alseodaphne Nees based on both morphological and molecular evidence. The combination of principal morphological characters to distinguish the two genera (Alseodaphnopsis vs. Alseodaphne) includes: 1) twigs thick, 4–11 mm in diameter, not obviously whitish in color vs. thin, 2.5–4.5 mm in diameter, obviously whitish in color; 2) terminal buds perulate vs. not perulate; 3) perianth lobes persistent at least in young fruit vs. early deciduous; 4) inflorescences relatively large, 8.5–35 cm long, generally many-flowered, with 3–4 order of branching vs. 3–20 cm long, few-flowered, with 1–2 orders of branching; and 5) mature fruit relatively large, 3–5 cm vs. < 2.5cm in diameter (Mo et al. 2017a). In addition, Alseodaphnopsis species are distributed in the northern marginal zone of Asian tropics in southwestern China (also in Hainan island) and northern Vietnam while Alseodaphne species are mostly found in the tropics of south and southeast Asia (Kostermans 1973; Li et al. 2008; Mo et al. 2017a).

During recent field surveys in southwestern China (Maguan, Yunnan Province) and northern Myanmar (Putao, Kachin State), two unknown Lauraceae species were collected. Based on both morphological and molecular evidence, they were confirmed as new species of Alseodaphnopsis and closely related to Alseodaphnopsis rugosa (Merr. & Chun) H. W. Li & J. Li and A. hainanensis (Merr.) H. W. Li & J. Li respectively. In the work of Mo et al. (2017b), the specimens collected in Maguan (Yunnan Province, China) were treated as synonyms of Alseodaphnopsis rugosa. In this paper, these new Alseodaphnopsis species are described and illustrated.

Material and methods

Morphological studies

Morphological characters of the two new Alseodaphnopsis species were examined in detail based on dried specimens and fresh materials in field observations and compared with possible relatives based on the specimens from the HITBC, IBK, IBSC and KUN herbaria as well as images of specimens available on JSTOR Global Plants (http://plants.jstor.org/).

Molecular studies and phylogenetic analyses

Total genomic DNA was extracted from silica-gel dried leaf material using the Plant Genomic DNA Kit (Tiangen, Beijing, China). Two nuclear DNA fragments, internal transcribed spacer (ITS) and the second intron of LEAFY gene (LEAFY intron II), were amplified and sequenced following the work of Li et al. (2011). ITS and LEAFY intron II sequences of other possibly related species involved here were obtained from GenBank according to the works of Li et al. (2011) and Mo et al. (2017a). Species examined in this study, voucher information, collection locality and GenBank accession numbers for ITS and LEAFY sequences are given in Table 1.

Species examined in this study, voucher information, collection locality 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 HQ697182 HQ697007
A. semecarpifolia Nees Arifiani DA658 (BO) Indonesia, Java HQ697184 HQ697015
Alseodaphnopsis (8)
A. andersonii (King ex Hook. f.) Kosterm. Li J. & Li L. 20070074 (HITBC) China, Yunnan FM957793 HQ697002
A. hainanensis Merr. Li L. & Wang Z. H. JFL07 (HITBC) China, Hainan MG188587 MG188634
Li L. & Wang Z. H. LMS10 (HITBC) China, Hainan MG188586 MG188633
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
A. petiolaris (Meissn.) Hook. f. 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 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 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. nanmu (Oliv.) Gamble 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

DNA sequences were aligned using Clustal X 2.1 (Larkin et al. 2007) and adjusted manually. The combined dataset including ITS and LEAFY intron II sequences for phylogenetic analysis was built according to the works of Li et al. (2011) and Mo et al. (2017a). 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 heuristic search was performed with 1000 random sequence addition replicates, tree-bisection-reconnection (TBR) swapping, MulTrees on, and all characters equally weighted. Bootstrap values of the internal nodes were obtained with 1000 bootstrap replicates. 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: TVM+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 Markov chain Monte Carlo (MCMC) algorithm was run for 10 million generations with a sampling frequency of one every 1000 generations, and the first 25% trees were discarded as burn-in.

Results

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. 1. All Alseodaphnopsis individuals investigated in the present study formed a monophyletic clade that receives a BS of 86% and a PP of 1.00. Within the Alseodaphnopsis clade, two well-supported subclades are found, each consisting of four species. The new species Alseodaphnopsis maguanensis is sister to A. rugosa (BS 89%, PP 1.00) and new species A. putaoensis is sister to A. hainanensis (BS 88%, PP 1.00).

Figure 1. 

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.

Taxonomic treatments

Alseodaphnopsis maguanensis L.Li & J.Li, sp. nov.

Figs 2, 3

Diagnosis

Alseodaphnopsis maguanensis is morphologically similar and phylogenetically closely related to A. rugosa, but can be distinguished by its much larger fruit (4–5 × 5–6 cm vs. ca. 2.5 × 3 cm), mature fruit color (brown vs. deep purple or black) and different fruiting phenology.

Type

China. Yunnan Province: Maguan County, Houcao, Gulinqing Provincial Natural Reserve, in tropical montane forest, 800 m a.s.l., 14 May 2016, flowering, Lang Li et al., GLQ26 (holotype: HITBC!).

Description

Trees evergreen, up to 20 m tall. Branchlets terete, 3–6 mm in diameter, grayish, glabrous, wrinkled, with lenticels and leaf scars. Terminal buds glabrous. Leaves clustered at apex of branchlet, alternate or subverticillate; petiole robust, 2–3 mm thick, 1.5–2.5 cm long, concave-convex; leaf blade green adaxially, glaucous abaxially when young but green or pale green when mature, oblong-obovate or oblong-oblanceolate, 12–32 × 3.5–9 cm, leathery, glabrous on both surfaces, midrib conspicuously elevated abaxially, impressed adaxially, lateral veins 8–12 pairs, veins and veinlets conspicuous, reticulate, elevated on both surfaces when dry, base cuneate, apex shortly acuminate. Panicles subterminal, clustered at apex of branchlet, 15–20 cm, many-flowered; peduncle 4.5–10 cm, glabrous. Pedicels slender, 5–8 mm, glabrous. Perianth lobes 6, glabrous outside, white pubescent inside, outer ones broadly ovate, ca. 2 × 1.5 mm, acute, inner ones broadly ovate, ca. 2.5 × 2 mm, acute, all deciduous when in fruit. Fertile stamens 9, ca. 2 mm in 1st and 2nd whorls, ca. 2.2 mm in 3rd whorl; filaments villous, almost as long as anthers in 1st and 2nd whorls, slightly longer than anthers in 3rd whorl, those of 3rd whorl each with 2 shortly stalked orbicular-cordate glands, others glandless; anthers of 1st and 2nd whorls ovate, with 2 upper smaller cells and 2 lower large cells, cells all introrse, anthers of 3rd whorl elliptic, with 4 extrorse cells. Staminodes conspicuous, ca. 1.5 mm, sagittate, stalked. Ovary ovoid, ca. 1.2 mm, glabrous, attenuate into a ca. 0.8 mm long style; stigma discoid, inconspicuous. Infructescence subterminal, 10–18 cm, robust, glabrous, with only one well-developed fruit. Fruit large, 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 cylindric, fleshy and warty when fresh.

Figure 2. 

Alseodaphnopsis maguanensis A habitat B branchlet with inflorescences C branchlet with mature fruit, immature fruit. Photographed by Lang Li.

Figure 3. 

Alseodaphnopsis maguanensis A outer perianth lobe (inside view) B inner perianth lobe (inside view) C third whorl stamen D pistil E flowering branch F second whorl stamen G staminode H first whorl stamen I flower (lateral view). Illustration by Ling Wang from Mo et al. (2017b).

Phenology

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

Etymology

The species is named after the type locality, Maguan County, in Yunnan Province, China.

Distribution and habitat

Currently known only from the type locality in Maguan, Yunnan Province, southwestern China. Tropical montane forests in valleys; ca. 800m.

Preliminary conservation status

Currently, A. maguanensis is only known from Maguan (Yunnan Province, China) with two populations, which are all located in a small natural reserve (ca. 71 km2), each population with less than 50 mature individuals (seedlings can be found near the mature individuals), and no other occurrence in adjacent regions of SE Yunnan and N Vietnam. Thus, the preliminary conservation status for A. maguanensis is suggested as critically endangered (CR C2a(i)) according to the IUCN Red List Categories and Criteria (IUCN 2012).

Additional specimen examined (paratype)

China. Yunnan Province: Maguan County, Shangba, Gulinqing Provincial Natural Reserve, in tropical montane forest, 800 m a.s.l., 28 August 2016, fruiting, Lang Li et al., 2016033 (HITBC!).

Alseodaphnopsis putaoensis L.Li, Y.H.Tan & J.Li, sp. nov.

Fig. 4

Diagnosis

Alseodaphnopsis putaoensis is morphologically similar to A. rugosa, but phylogenetically closely related to A. hainanensis. It can be distinguished from them by its fruit stalk characters (apex slightly dilated, not fleshy, red and warty when fresh vs. apex dilated, nearly cylindric, fleshy, red and warty when fresh), much larger fruit (6–6.5 × 7–10 cm vs. ca. 2.5 × 3 cm and 1.2–2 cm), mature fruit color (brown vs. deep purple or black) and different fruiting phenology.

Type

Myanmar. Kachin State: Putao County, on the way from Masabu to Namti, in tropical montane forest, 1000 m a.s.l., 13 May 2017, fruiting, Lang Li & Hui Ma MM271 (holotype: HITBC!).

Description

Trees evergreen, up to 15 m tall. Branchlets terete, robust, 7–10 mm in diameter, brownish or dark brown, glabrous, wrinkled, with lenticels and leaf scars. Terminal buds glabrous. Leaves clustered at apex of branchlet, subverticillate; petiole robust, 2–3 mm thick, 2.5–4.5 cm long, concave-convex; leaf blade green adaxially, glaucous abaxially, oblong-obovate or oblong-oblanceolate, 18–35 × 6–9 cm, leathery, glabrous on both surfaces, midrib conspicuously elevated abaxially, impressed adaxially, lateral veins 8–12 pairs, veins and veinlets conspicuous, reticulate, elevated on both surfaces when dry, base cuneate, apex shortly acuminate. Flowers unknown. Infructescence subterminal, 8–10 cm, robust, glabrous, with only one well-developed fruit. Fruit large, oblate, 6–6.5 × 7–10 cm, immature fruit green, brown when mature, fruit stalk robust, 3–4 mm in diameter, apex slightly dilated, 5–6 mm in diameter.

Figure 4. 

Alseodaphnopsis putaoensis A habitat B branchlet with immature fruit C branchlet with immature fruit, immature and mature fruits. Photographed by Lang Li.

Phenology

Individuals with immature or mature fruits have been collected in May, fruiting may be from April to June.

Etymology

The species is named after the type locality, Putao County, in Kachin State, Myanmar.

Distribution and habitat

Currently known only from the type locality in Putao, Kachin State, northern Myanmar. Tropical montane forests on mountain slopes or in valleys; 600–1400m.

Preliminary conservation status

During the field survey in Putao (Kachin State, Myanmar), several populations of A. putaoensis were found and at least two of them with more than 50 mature individuals (seedlings could be found near the mature individuals) each. In future field surveys, potential populations and more individuals are expected to be found in Putao and adjacent regions. Currently, some localities of A. putaoensis have not been legally protected. The habitat fragmentation, as well as ongoing road construction and continuous logging, are threatening its survival. Thus, the preliminary conservation status for A. putaoensis is suggested as vulnerable (VU C12a(i)) according to the IUCN Red List Categories and Criteria (IUCN 2012).

Additional specimen examined (paratype)

Myanmar. Kachin State: Putao County, on the way from Masabu to Namti, in tropical montane forest, 900 m a.s.l., 13 May 2017, fruiting, Lang Li & Hui Ma MM254 (HITBC!).

Discussion

The close relationships of Alseodaphnopsis maguanensis, A. rugosa, A. putaoensis and A. hainanensis were indicated by the phylogenetic analyses. They formed a well-supported subclade within the Alseodaphnopsis clade, and A. maguanensis is sister to A. rugosa while A. putaoensis is sister to A. hainanensis (Fig. 1). Alseodaphnopsis maguanensis, A. putaoensis and A. rugosa are very similar in their vegetative characters (e.g. leaf and branchlet characters), but A. hainanensis can be easily distinguished from them by its narrowly elliptic and smaller leaves (vs. oblong-obovate or oblong-oblanceolate and larger leaves of A. maguanensis, A. putaoensis and A. rugosa) (Li et al. 2008). The fruit stalk and fruit characters are very important to distinguish Alseodaphnopsis maguanensis, A. putaoensis and A. rugosa. Alseodaphnopsis putaoensis has the largest fruit (6–6.5 × 7–10 cm, brown when mature) with the fruit stalk slightly dilated at the top (not fleshy and warty when fresh), while A. maguanensis and A. rugosa have relative smaller fruits (4–5 × 5–6 cm, brown when mature and ca. 2.5 × 3 cm, deep purple or black when mature) with the fruit stalks distinctly dilated at the top (nearly cylindric, fleshy and warty when fresh).

Alseodaphnopsis maguanensis, A. rugosa, A. putaoensis and A. hainanensis also have different fruiting phenologies. Alseodaphnopsis maguanensis is fruiting from July to September, both immature and mature fruits can be found in August. According to the work of Li et al. (2008), Alseodaphnopsis rugosa is fruiting from July to December. However, based on specimen and field observations, very few individuals with young fruits were collected in August, most fruiting individuals were collected from October to December. For Alseodaphnopsis putaoensis, individuals with immature or mature fruits have been collected in May, fruiting may be from April to June. Alseodaphnopsis hainanensis is fruiting from October to February (Li et al. 2008). A detailed comparison of the morphological differences among these four taxa, as well as their phenologies and distributions, is given in Table 2. Unfortunately, the flowers of Alseodaphnopsis putaoensis and A. rugosa remain unknown. The evidence from both phylogenetic and morphological analyses support the recognition of Alseodaphnopsis maguanensis and A. putaoensis as distinct species in the genus.

Comparison of key morphological characters, phenologies and distributions of Alseodaphnopsis maguanensis, A. rugosa, A. putaoensis and A. hainanensis.

Morphological characters Alseodaphnopsis maguanensis Alseodaphnopsis rugosa Alseodaphnopsis putaoensis Alseodaphnopsis hainanensis
Leaf blade oblong-obovate or oblong-oblanceolate, 12–32 × 3.5–9 cm, green adaxially, glaucous abaxially when young but green or pale green when mature oblong-obovate or oblong-oblanceolate, 15–36 × 4–10 cm, green adaxially, glaucous abaxially oblong-obovate or oblong-oblanceolate, 18–33 × 6–9 cm, green adaxially, glaucous abaxially narrowly elliptic, 6–16 × 1.5–4.2 cm, green adaxially, glaucous abaxially when young but green or pale green when mature
Infructescence with only one well-developed fruit with one or several well-developed fruits with only one well-developed fruit with one or several well-developed fruits
Fruit stalk apex dilated, 5–10 mm in diam., sometimes nearly cylindric, fleshy and warty when fresh apex dilated, nearly cylindric, 5–8 mm in diam., fleshy, red and warty when fresh apex slightly dilated, 5–6 mm in diam. apex dilated, nearly cylindric, 5–8 mm in diam., fleshy, red and warty when fresh
Fruit oblate, 4–5 × 5–6 cm, brown when mature oblate, ca. 2.5 × 3 cm, deep purple or black when mature oblate, 6–6.5 × 7–10 cm, brown when mature globose or ovoid, 1.2–2 cm, deep purple or black when mature
Flowering Phenology May–Jun Jul
Fruiting Phenology Jul–Sep Jul–Dec (Fruits mostly found from Oct to Dec) Apr–Jun Oct–Feb of next year
Distribution SW China (Yunnan) S China (Hainan) N Myanmar (Kachin) S China (Hainan), N Vietnam (Lào Cai)

With Alseodaphnopsis maguanensis, A. putaoensis and the recently described A. ximengensis H. W. Li & J. Li from Ximeng, Yunnan Province, China (Mo et al. 2017a) included, Alseodaphnopsis currently has only 11 recorded species, indicating that the species diversity of the genus is still in need of investigation and open to discovery. More new species of Alseodaphnopsis are expected to be discovered in the northern marginal zone of the Asian tropics when more field investigations are conducted in this region.

Acknowledgements

The authors are grateful to the staff of Gulinqing Provincial Nature Reserve, Maguan, Yunnan Province, China and Hkakaborazi National Park, Putao, Kachin State, Myanmar for their help during the field survey. This work was financially supported by grants from Southeast Asia Biodiversity Research Institute, Chinese Academy of Sciences (Y4ZK111B01) and Yunnan Applied Basic Research Projects (2017FB033).

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