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Research Article
Mallotus bullatus (Euphorbiaceae), a new species from Southwest China based on morphological characters and phylogenetic evidence
expand article infoJiang-Hong Yu‡, Zheng-Ren Chen§, Ming-Tai An‡, Deng-Li Yu§, Feng Liu‡, Jian Xu|, Yu-Bin Tang‡, Yi-Ran Wang‡, Hua-Kai Zou‡
‡ Guizhou University, Guiyang, China
§ Guizhou Maolan National Nature Reserve Administration, Libo, China
| Guizhou Botanical Garden, Guiyang, China

Corresponding author: Ming-Tai An ( gdanmingtai@126.com )

Academic editor: Geoffrey Levin
© 2024 Jiang-Hong Yu, Zheng-Ren Chen, Ming-Tai An, Deng-Li Yu, Feng Liu, Jian Xu, Yu-Bin Tang, Yi-Ran Wang, Hua-Kai Zou.
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: Yu J-H, Chen Z-R, An M-T, Yu D-L, Liu F, Xu J, Tang Y-B, Wang Y-R, Zou H-K (2024) Mallotus bullatus (Euphorbiaceae), a new species from Southwest China based on morphological characters and phylogenetic evidence. PhytoKeys 249: 13-25. https://doi.org/10.3897/phytokeys.249.131824
Open Access

Abstract

Mallotus bullatus M.T.An & J.H.Yu, sp. nov. (Euphorbiaceae), a species new to science discovered in Guizhou, China, is described and illustrated here, and its phylogenetic position among other Mallotus species is presented. Morphological, micro-morphological, and molecular evidence is presented as attestation of its novelty. The new species morphologically resembles M. philippensis var. reticulatus and M. philippensis var. philippensis, but it clearly differs by having bullate leaf surfaces (vs. not bullate), leaf margins entire or nearly so (vs. entire or nearly so in M. philippensis var. philippensis and coarsely serrate in M. philippensis var. reticulatus), leaf margins sometimes bearing red glands (vs. red glands absent), 5 sepals in staminate flowers (vs. 3–4 in M. philippensis var. philippensis and 4 in M. philippensis var. reticulatus), fruits with spines (vs. spines absent in M. philippensis var. philippensis and present in M. philippensis var. reticulatus), and abaxial leaf epidermal scattered and clustered vein hairs 0.1–0.8 mm long (vs. 0.04–0.28 mm long in M. philippensis var. philippensis and 0.05–0.1 mm long in M. philippensis var. reticulatus). Molecular phylogenetic analysis (BS = 100% / BS = 96%, PP = 1 / PP = 1) provides strong evidence supporting M. bullatus as a new species within the genus Mallotus and supports its placement in M. sect. Philippinenses as sister to M. philippensis.

Key words

Euphorbiaceae, Guizhou province, karst, molecular identification

Introduction

Mallotus Lour. (Euphorbiaceae) is a large genus comprising approximately 150 species (Sierra et al. 2005), predominantly consisting of shrubs or trees, seldom climbers. It is mainly distributed in tropical and subtropical regions of Asia, Australia, and the Pacific, with a few species found in tropical Africa and Madagascar (Kulju et al. 2007a; Sierra et al. 2007). In China, there are approximately 30 species of Mallotus, mainly distributed in southern provinces and regions. The bark of some species is used for making ropes, and the seed oil is used for soap and industrial oils (Liao and Liu 1958). Additionally, Mallotus species are important medicinal plants and sources of dye (Kumar et al. 2006; Sharma and Varma 2011; Dhaker and Sharma 2014). The genus Mallotus is an important component of forest vegetation (Slik et al. 2003; Eichhorn 2006), exhibiting a variety of life history strategies. Some species act as early successional pioneers, while others are climax species. The genus occurs in a wide range of habitats at low elevations (Sierra et al. 2007).

The genus Mallotus was established by De Loureiro in 1790 based on Mallotus cochinchinensis Lour. (Loureiro 1790). In the latest taxonomic studies of Euphorbiaceae (Webster 1994; Radcliffe-Smith 2001), the genus Mallotus is classified in the subtribe Rottlerinae Meisn. In addition, Mallotus has morphological, distribution and ecological similarities with Macaranga Thouars, another large genus in the Euphorbiaceae. Two phylogenetic studies (Slik and van Welzen 2001; Kulju et al. 2007b) specifically investigated the relationships of Mallotus with related genera. Kulju et al. (2007b) identified three clades, with the majority of Mallotus (Mallotus sensu stricto [s.str.]) forming a sister group to several small genera within the Macaranga clade. Sierra et al. (2010) compiled various datasets including plastid (matK) and nuclear (ITS) DNA sequences, macro-morphological characteristics, and leaf anatomical data, providing a detailed analysis of the phylogeny of Mallotus. The study revealed that Mallotus sect. Mallotus, sect. Polyadenii Pax & K.Hoffm., and sect. Stylanthus Pax & K.Hoffm. are monophyletic, while sect. Axenfeldia (Baill.) Pax & K.Hoffm. and sect. Rottleropsis Müll.Arg. are polyphyletic, and sect. Philippinenses Pax & K.Hoffm. is paraphyletic.

In 2023, during a botanical survey in the Maolan National Nature Reserve in Guizhou, China, we discovered a possibly new species of Euphorbiaceae. After more than a year of field investigations and specimen collection (GZAC-MU-0001), we conducted a field investigation on new species in Maolan National Nature Reserve, Guizhou Province, including photographing its characteristics and collecting seven live specimens. We found that its morphological characteristics resemble those of the genus Mallotus. To effectively distinguish the new species from other Mallotus species, this study utilized morphology, including pollen and leaf epidermal micromorphology, and molecular phylogenetics using ITS and matK sequences. The results led to the conclusion that the putative new species represents a new taxon.

Materials and methods

Morphology

Morphological features of leaves, inflorescences, flowers, and capsules were carefully observed and measured in the field, followed by detailed examination in the laboratory. Additionally, we compared specimens based on field observations and photographs taken, as well as studied related species using FAA-fixed materials and dried specimens (GZAC).

Leaf epidermis and pollen grains

The mature, complete pollen grains and leaves collected from the field were used to measure characters through a dissecting microscope. Subsequently, they were mounted on stubs with double-sided tape, coated with a layer of gold, and then photographed using a Hitachi S-4800 scanning electron microscope. The micro-morphological characteristics of the pollen grains are described according to Wang and Wang (1983) and Nowicke and Takahashi (2002). The average size of the pollen grains is calculated based on 20 samples. The micromorphological features of the leaf epidermis of the genus Mallotus are stable genetic characteristics that show certain interspecific differences, reflect certain phylogenetic relationships, and can provide a basis for classification and species identification within the genus (Raju and Rao 1977; Alyas et al. 2020). Therefore, this study also investigates the leaf epidermal micromorphology of Mallotus philippensis H.Karst., including both var. philippensis and var. reticulatus (Dunn) F.P.Metcalf. The description of leaf micromorphological features follows Fiala et al. (1994) and Živa et al. (2012), and based on seven specimens collected in the field, including the holotype and the two paratypes.

Taxon sampling and DNA sequencing

We used a total of 36 species of Mallotus (Euphorbiaceae) in this study, including two individuals of the new species, and one outgroup species: Macaranga trichocarpa (Zoll.) Müll.Arg. We chose to use two molecular markers: ITS (ITS-1, 5.8S, and ITS-2) and matK. The ITS sequence, a highly reiterated tandem sequence in the nuclear genome, exhibits rapid changes, providing abundant variation and informative sites (Nürk et al. 2015) and a high level of species resolution accuracy (Chinese Plant Bol Group et al. 2011). The matK gene is one of the fastest-evolving genes in the chloroplast genome. It is easy to align and widely used in the study of families, genera and species (Khidir and Hongping 1997).

We extracted DNA sequences from fresh leaves of the new species and M. philippensis var. reticulatus using a modified CTAB protocol from Doyle and Doyle (1987), followed by PCR amplification and sequencing following the protocols in referred the methods of White et al. (1990) and Taberlet et al. (1991). We downloaded DNA sequences from GenBank for the two molecular markers for the remaining species used. Taxa and GenBank accession numbers are listed in Suppl. material 1: table S1.

Phylogenetic analysis

Sequences were aligned using default parameters in Clustal X v.1.83 (Thompson et al. 1997), followed by manual adjustments in BioEdit v.7.0 (Hall 1999). The phylogenies were constructed using Maximum Likelihood (ML) as implemented in PhyloSuite (Zhang et al. 2020) and Bayesian Inference (BI) as implemented in MrBayes v.3.0b4 (Ronquist and Huelsenbeck 2003), with the ITS and matK sequences analyzed separately. For the the nucleotide substitution model was chosen using the Akaike Information Criterion (AIC) in Modeltest v.3.06 (Posada and Crandall 1998), with the GTR+I+G model chosen for ITS and the GTR+I model for matK. For the ML analyses, the nucleotide substitution model was chosen using AIC in ModelFinder (in PhyloSuite), with the GTR+G model chosen for both regions.

Results

Taxonomic treatment

Mallotus bullatus M.T.An & J.H.Yu, sp. nov.

urn:lsid:ipni.org:names:77351570-1
Fig. 1

Type

China. • Guizhou Province, Libo County, Dawn township aquatic animals, 25°19'N, 107°56'E, alt. 700 m, 29 April 2024, Ming-tai An, Jiang-hong Yu, Jian Xu, Feng Liu GZAC-MU-001 (holotype GZAC!).

Figure 1. 

Mallotus bullatus A habitat B, C plant D leaf from above E leaf from below F, G leaf lower surface H–J inflorescences K pistillate flower L, M staminate flower sepals N infructescence with mature capsules O capsule with part removed to show seed P capsule in transverse section.

Diagnosis

M. bullatus can be distinguished from M. philippensis var. reticulatus and M. philippensis var. philippensis by having leaves with bullate surfaces and entire or nearly so margins, sometimes bearing red glands, 5 sepals in the staminate flowers, fruits with spines, pollen grains tricolporate with obvious furrows containing protrusions in the apertures, and abaxial leaf scattered and clustered vein hairs 0.1–0.8 mm long (Table 1).

Table 1.
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Comparison of characteristic of three species of M. bullatus, M. philippensis var. philippensis, and M. philippensis var. reticulatus.

Character M. philippensis var. philippensis M. philippensis var. reticulatus M. bullatus
Habit Small tree or shrub Shrub Shrub
Upper leaf midrib hairys No Yes No
Leaf margins Entire or nearly so Coarsely serrate Entire or nearly so
Leaf margins bearing red glands No No Sometimes
Leaf surface bullate No No Yes
Length of solitary or clustered hairs on leaf abaxial veins 0.04-0.28 mm (Zhang, 2018) 0.05-0.1 mm 0.1-0.8 mm
Number of staminate sepals 3-4 4 5
Pollen size 15.5(17.5)-(11.6)15.5 μm (Nowicki and Takahashi 2002) 19 × 20 μm 22 × 20 μm
Fruits with spines No Yes Yes

Description

Shrubs , 1.5–2.5 m tall; twigs, young leaves, and inflorescences densely covered with yellowish-brown disc-shaped glandular hairs. Leaves simple, alternate, ovate or lanceolate, 5–18 (-22) × 3–6 cm, thickly papery, apex acuminate, base rounded or cuneate, margins entire or nearly so, sometimes bearing red glands, surface bullate, upper surface glabrous, lower surface densely grayish-yellow clustered-tomentose, with long soft solitary or clustered hairs on the veins, and scattered red disc-like glands; basal veins 3, lateral veins 3–4 pairs, looped and joined near the margin; extrafloral nectaries, 2–4, brown, near the base; petiole round 2–5 (-9) cm long, slightly pulvinate at both ends, covered with clustered hairs. Inflorescences racemose, terminal, solitary or clustered, solely staminate or pistillate, or mixed with pistillate flowers in lower part and staminate ones in upper part; sometimes apparently bisexual flowers also present. Staminate inflorescences 5–10 cm long, bracts ovate, ca. 1 mm long, pedicel 1–2 mm long, calyx lobes 5, oblong, ca. 2 mm long, densely covered with stellate hairs, with red disc-like glands; stamens 28–30. Pistillate inflorescences s 3–8 cm long, bracts ovate, about 1 mm long; pedicels ca. 1–2 mm long; calyx lobes 4, ovate, densely covered with stellate hairs outside, ca. 3 mm long; ovary hairy, stigmas 3 split, 3–4 mm long, stigmas densely set with feather-like papillae on upper surface; some pistillate flowers sometimes bisexual, then with 1 or 2 stamens, the filaments almost as long as the anthers. Bisexual inflorescences 5–10 cm long, with 3–6 staminate flowers at the apex, lower part entirely pistillate; bracts ovate. Capsule subglobose, with spines, ca. 6–8 mm in diameter, fruit wall thickness ca.1–2 mm, 3 carpellate, densely covered with red disc-like glands; seeds black, ovate or globose, naked with late mature stage.

Distribution and habitat

This species is known only from the karst landscape of Libo County, Guizhou Province, China (Fig. 2), alt. 700–900 m.

Figure 2. 

Geographical distribution of Mallotus bullatus.

Phenology

Flowering from April to May, and fruiting from May to August.

Etymology

“Bullatus” specifically refers to the convex leaf areoles.

Local name

Simplified Chinese: 荔波野桐; Chinese Pinyin: lì bō yě tóng.

Leaf epidermis and palynology

Pollen grains of M. bullatus are spheroidal, with a size of 22 × 20 μm, L(long)/W(width) = 1.1, and tricolporate. They feature tricolporate furrows containing protrusions (Fig. 3A–C). Pollen grains of M. philippensis var. reticulatus are also spheroidal, 19 × 20 μm, L/W ratio 0.95, without distinct furrows (Fig. 3D–F). The lower epidermis of M. bullatus leaves bears evenly distributed elliptical glands measuring 100 × 70 μm (E1 × E2: length of long equatorial axis × length of short equatorial axis), is densely covered with short clustered hairs, and has long (0.1–0.8 mm long) solitary or clustered hairs on the veins (Fig. 3G, H). Similarly, the lower epidermis of M. philippensis var. reticulatus exhibits elliptical glands measuring 80 × 70 μm (E1 × E2), is densely covered with short clustered hairs, and has solitary or clustered hairs 0.05–0.15 mm long on the veins (Fig. 3J, K). The upper epidermis of both M. bullatus and M. philippensis var. reticulatus leaves is smooth (Fig. 3I, L).

Figure 3. 

Scanning electron microscope images of Mallotus leaf epidermis and pollen grains A–C pollen grains of M. bullatus D–F pollen grains of M. philippensis var. reticulatus G–I lower leaf epidermis of M. bullatus J–L lower leaf epidermis of M. philippensis var. reticulatus.

Conservation status

During the period of 2023–2024, we sampled the population of M. bullatus and discovered two additional distribution points near the species initial discovery location (Fig. 2). Each site contained approximately 30 plants. The habitat of M. bullatus mainly occurs in karst scrublands, distributed from the foothills to the middle of the mountains. The plant habitat features poor soil fertility, low water retention capacity, and frequent drought conditions. Due to our current insufficient comprehensive assessment of the survival status and threats to M. bullatus, we cannot provide specific distribution information about this population. Therefore, we recommend categorizing M. bullatus as Data Deficient “DD” (IUCN 2022).

Morphological comparisons

Morphologically, the new species is similar to M. philippensis in having alternate leaves, basal veins 3, and racemose inflorescences. However, the new species can be distinguished from M. philippensis by its bullate leaf surface (vs. not bullate), leaf margins entire or nearly so (vs. entire or nearly so in M. philippensis var. philippensis, and coarsely serrate in M. philippensis var. reticulatus), leaf margins sometimes bearing red glands (vs. not red glands), fruits with spines (vs. absent in M. philippensis var. philippensis and present in M. philippensis var. reticulatus), 5 sepals in staminate flower (vs. 3–4 in M. philippensis var. philippensis and 4 in M. philippensis var. reticulatus) (Table 1, Suppl. material 1: fig. S1).

Phylogenetic position

Nuclear data phylogenetic analyses

The length of the aligned ITS sequences of M. bullatus is 760 bp. Based on a dataset of 28 ITS sequences with 182 informative loci, phylogenetic relationships were analyzed using both Bayesian Inference (BI) and Maximum Likelihood (ML) methods (Fig. 4). The two sequences from the new species both originate from the same population, forming a strongly supported monophyletic clade (Fig. 4: BS = 100%, PP = 1). Mallotus philippensis var. philippensis and M. philippensis var. reticulatus are sister taxa with strong support (Fig. 4: BS = 100%, PP = 1), and they form a strongly supported sister group relationship with M. bullatus (Fig. 4: BS = 98%, PP = 1).

Figure 4. 

Partial Bayesian consensus phylogram based on ITS sequences. Numbers above branches are Bayesian posterior probabilities (PP) and Bootstrap probabilities (BS) (only PP values > 0.70, BS > 80 shown).

Plastid data phylogenetic analysis

The aligned matK sequences of M. bullatus are 2000 bp in length. Based on a dataset of 27 matK sequences with 557 informative loci, both Bayesian Inference (BI) and Maximum Likelihood (ML) analyses indicate that the two sequences from the new species form a strongly supported monophyletic clade (Fig. 5: BS = 96%, PP = 1). Mallotus philippensis var. philippensis and M. philippensis var. reticulatus forms a weakly supported sister clade (Fig. 5: BS = 80%, PP = 0.82), and they form a strongly supported sister group relationship with M. bullatus (Fig. 5: BS = 100%, PP = 1).

Figure 5. 

Partial Bayesian consensus phylogram based on matK sequences. Numbers above branches are Bayesian posterior probabilities (PP) and Bootstrap probabilities (BS) (only PP values > 0.70, BS > 80 shown).

Discussion

According to the classification in Sierra et al. (2010), M. bullatus belongs to sect. Philippinenses, characterized by shrubby habit, alternate leaves, 3 basal leaf veins, and capsules densely covered with orange-red glands. Both phylogenetic trees (ITS and matK; Figs 4, 5) indicate that M. bullatus is a distinct member of the genus, and furthermore, support its sister group relationship with M. philippensis var. philippensis plus, M. philippensis var. reticulatus; these three species form a weakly supported clade with M. repandus (Figs 4, 5), also in sect. Philippinensis, thus corroborating the evidence provided by the morphological and micro-morphological observations. The two known populations of M. bullatus show no consistent morphological differences. Although M. bullatus forms a clade with M. philippensis var. philippensis and M. philippensis var. reticulatus, it differs from both taxa in its bullate leaf surfaces, length of hairs on the leaf abaxial veins, and number of sepals in the staminate flowers (Table 1).

Additional specimen examined

Mallotus philippensis var. reticulatus (Dunn) F. P. Metcalf. —China. Fujian: Collected on Mr. Dunn’s expedition to Central, China. Apr. to Jun., 1905, 3429 (HH); West lake, Chenxiang town, Changtai district, Zhangzhou city. Jun. 11, 1976, Wang QJ, 012320 (AU). Jangxi: Yangling, Chongyi County, Ganzhou City, 24°29'N, 103°54'E alt. 1092 m, May 15, 2024, Yu JH, Tang YB, Wang YR, 20240501 (GZAC). M. philippensis var. philippensis (Lam.) Müll. Arg. — Yunan: roadside at the edge of forests, alt. 800 m. 2000, Shui YM, Chen WH 13773 (PE). Sichuan: Jinjia Village, Sutie National Nature Reserve, Panzhihua, Sichuan Province, 26°37'29.7"N, 101°33'03.1"E, alt. 1653 m. Sept. 16, 2010. Yang Y, Huang JH, Yang YQ, Liu B, Ye JF 551 (PE). Paratypes. Mallotus bullatus — Guizhou: Rao gu village, Dawn township aquatic animals, Libo County, 25°19'N, 107°56'E, alt. 800 m, Apr. 29, 2024, An MT, Yu JH, Xu J, Liu F 202306-1 (GZAC); La nei village, Libo County, 28°21'N, 107°56'E, alt. 750 m, Apr. 30, 2024, An MT, Yu JH, Xu J, Liu F 202306-2 (GZAC). Other specimens. Mallotus bullatus — Guizhou: Yiba Mountain, Lane Village, Dawn township aquatic animals, Libo County, 25°16'N, 107°55'E, alt. 950 m, Jul. 26, 2024, Yu JH, Tang YB, Liu F 001 (GZAC); Yiba Mountain, Lane Village, Dawn township aquatic animals, Libo County, 25°16'N, 107°55'E, alt. 780 m, Jul. 28, 2024, Yu JH, Tang YB, Liu F 002 (GZAC); Dawn township aquatic animals, Libo County, 25°16'N, 107°57'E, alt. 840 m, Jul. 22, 2024, Yu JH, Tang YB, Liu F 003 (GZAC); Dawn township aquatic animals, Libo County, 25°16'N, 107°57'E, alt. 700 m, Jul. 22, 2024, Yu JH, Tang YB, Liu F 004 (GZAC).

Acknowledgements

The authors would like to thank Hong-Fen Hu and Xu Wu for their support in the sequencing process and photographing plants, as well as Yi-Fei Xie and Xiang-Dong Qiu for their help in collecting samples of Mallotus philippensis var. reticulatus in Jiangxi Province. Thank you to Geoffrey Levin, Peter van Welzen and an anonymous reviewer for providing input to improve 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 supported by the “Investigation and Monitoring Project of Maolan Large-scale Dynamic Plot of Karst Forest Eco-system in South China (2023–23), “Survey and Assessment of Newly Added National Key Protected Wild Plant Resources in Guizhou Province (Three stage) (MCHC–ZD20242057)” and “Research on the diversity of Corybas fanjingshanensis mycorrhizal fungi of rare and endangered orchids endemic in Guizhou (QKHJC [2023]1Y235)”.

Author contributions

Jiang Hong Yu and Zheng Ren Chen completed all the work of this paper together, they contributed equally to this work and are the first co- authors of this paper. MingTai An planned and guided the writing of the whole paper, participated in the field investigation and identification work, acted as the corresponding author of the paper. Deng Li Yu participated in the field investigation and completed part of the data processing and content writing and guide the writing of the paper. Feng Liu Collect plant specimens in the field and take photographs for identification. Jian Xu participated in field investigation and compared plant specimens to identify species. Yu Bin Tang participated in field investigation and compared plant specimens to identify species. Yi Ran Wang participated in field investigation and data collation. Hua Kai Zou participated in field investigation and content writing.

Author ORCIDs

Jiang-Hong Yu https://orcid.org/0000-0003-1765-8557

Ming-Tai An https://orcid.org/0000-0003-3886-0287

Deng-Li Yu https://orcid.org/0009-0004-4577-6678

Feng Liu https://orcid.org/0009-0003-7031-4494

Jian Xu https://orcid.org/0000-0002-0714-0917

Yu-Bin Tang https://orcid.org/0009-0007-8460-2242

Yi-Ran Wang https://orcid.org/0009-0008-6030-8601

Hua-Kai Zou https://orcid.org/0009-0009-2029-6068

Data availability

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

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Supplementary material

Supplementary material 1 

Additional information

Jiang-Hong Yu, Zheng-Ren Chen, Ming-Tai An, Deng-Li Yu, Feng Liu, Jian Xu, Yu-Bin Tang, Yi-Ran Wang, Hua-Kai Zou

Data type: docx

Explanation note: table S1. Taxa and GenBank accession numbers for thesequences used in this study. fig. S1. Mallotus philippensis var. philippensis (Lam.) Müll.Arg. and M. philippensis var. reticulatus (Dunn) F.P.Metcalf. A, B leaf of Mallotus philippensis var. reticulatus C holotype of Mallotus philippensis var. philippensis D holotype of Mallotus philippensis var. reticulatus E merotype of Mallotus philippensis var. reticulatus.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
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