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Nghiên Cứu Khả Năng Kết Hợp Và Mức Độ Chống Chiu Sâu ISSN: 1859-2171 TNU Journal of Science and Technology 207(14): 33 - 40 e-ISSN: 2615-9562 TAXONOMY AND PHYLOGENY OF VIETNAMESE SPONDIAS L. (ANACARDIACEAE) Chi Toan Le1*, Van Du Nguyen2,3, Wyckliffe Omondi Omollo4, Bing Liu4 1Ha Noi Pedagogical University No. 2, 2Institute of Ecology and Biological Resources - Vietnam Academy of Science and Technology, 3Graduated University of Vietnam Academy of Science and Technology, 4Institute of Botany, Chinese Academy of Sciences, ABSTRACT Spondias is a small genus of Anacardiaceae. Vietnamese Spondias has remained taxonomically unresolved and the infrageneric relationship within the genus has been disputed. Here, we present molecular phylogenetic analyses of this genus and its close relatives using a combined dataset of the chloroplast rbcL, matK, and trnL-F regions. Molecular analyses strongly supported the monophyly of Spondias with two major clades and Vietnamese Spondias was placed within Asian Spondias clade. Based on both morphological and molecular data, we recognized two species of Spondias: S. dulcis Parkinson and S. pinnata (L. f.) Kurz in Vietnam. The key and description for Vietnamese Spondias species were provided. We also suggested to recognize Spondias petelotii as a synonym of Allospondias lakonensis. Keywords: Molecular; Taxonomy; Phylogeny; Spondias; Synonym; Anacardiaceae Ngày nhận bài: 08/7/2019; Ngày hoàn thiện: 07/8/2019; Ngày đăng: 09/9/2019 NGHIÊN CỨU PHÂN LOẠI VÀ PHÁT SINH LOÀI CỦA CHI CÓC SPONDIAS L. (ANACARDIACEAE) Ở VIỆT NAM Lê Chí Toàn1*, Nguyễn Văn Dư2,3, Omollo Omondi Wyckliffe4, Liu Bing4 1Trường Đại học Sư phạm Hà Nội 2, 2Viện Sinh thái Tài nguyên Sinh vật - Viện Hàn lâm Khoa học Việt Nam, 3Học viện Khoa học và Công nghệ - Viện Hàn lâm Khoa học Việt Nam, 4Viện Thực vật học - Viện Hàn lâm Khoa học Trung Quốc TÓM TẮT Chi Cóc (Spondias L.) là một chi nhỏ của họ Xoài. Việc sắp xếp phân loại và tìm hiểu mối quan hệ di truyền của chi Cóc ở Việt Nam là chưa rõ ràng và còn tồn tại một số vấn đề. Nghiên cứu này tiến hành phân tích mối quan hệ phát sinh loài của chi Cóc và họ hàng gần gũi của chi này dựa trên dữ liệu sinh học phân tử là các đoạn gen lục lạp rbcL, matK, và trnL-F. Kết quả phân tích dữ liệu phân tử ủng hộ mạnh mẽ rằng chi Cóc là chi đơn phát sinh với hai nhánh phát sinh chính là nhánh Cóc Nam Mỹ và nhánh Cóc châu Á; Cóc Việt Nam nằm trong nhánh Cóc châu Á. Dựa trên cả dữ liệu phân tử và hình thái, nghiên cứu này ghi nhận Cóc Việt Nam bao gồm hai loài: Spondias dulcis Parkinson và Spondias pinnata (L. f.) Kurz. Khóa định loại và mô tả cho các loài Cóc Việt Nam được cung cấp. Nghiên cứu này cũng chỉ ra rằng Spondias petelotii là đồng nghĩa của Allospondias lakonensis. Từ khóa: Phân tử; Phân loại; Phát sinh loài; Spondias; Đồng nghĩa; Anacardiaceae Received: 08/7/2019; Revised: 07/8/2019; Published: 09/9/2019 * Corresponding author. Email: [email protected] http://jst.tnu.edu.vn; Email: [email protected] 33 Lê Chí Toàn và Đtg Tạp chí KHOA HỌC & CÔNG NGHỆ ĐHTN 207(14): 33 - 40 1. Introduction philippinensis is similar to the genus Spondias L., the type genus of the subfamily Solenocarpus such as: eucamptodromous Spondiadoideae Kunth ex Arn., is a small secondary venation, single narrowly flabellate genus of fruit trees of Anacardiaceae with 18 style, single stigma, unicarpellate ovary, species [1]. Members of Spondias are mainly strongly oblique fruit. Futhermore, S. distributed in tropical Asia, America and one philippinensis and Solenocarpus indicus species in Madagascar. The Spondias species Wight & Arn. morphologically share floral show significant economic importance with features such as: apert calyx, valvate corolla, various fruits that are used both as human and single narrowly flabellate style. Thus, animal food [2]. Spondias philippinensis should be kept out of Spondias and placed in Solenocarpus. In The taxonomic history of Spondias was quite addition, Haplospondias brandisiana (Kurz) complex. Spondias was one of the first genera Kosterm. was considered as distinct from of Anacardiaceae described by Linnaeus with Spondias based on simple leaves without an the type species S. mombin L. published in intramarginal vein and a single style with an 1753 [1]. Bentham & Hooker (1862) [3] oblique stigma [1]. divided the family Anacardiaceae into two tribes, the Anacardieae and Spondieae. Michell & Daly (2015) [1] also disscused to Subsequently, Marchand (1869) [4] published Solenocarpus indicus and Spondias dulcis. the tribe Spondiadeae (as Spondieae) and was They emphasized that the placement of the first to formulate a relatively modern Spondias dulcis in Evia is not correct, this concept of Spondias, in which he included species should be treated as a member of Evia Blume, Cytheraea Wight & Arn. and Spondias based on both morphologycal and Wirtgenia Jung. ex Hassk. On the other hand, molecular data; while, Solenocarpus indicus of the taxa he either accepted in Spondias or Wight & Arn. should be separated from recognized as synonyms of species in the Spondias [1]. Additionaly, the situations of genus, four are considered here to belong to Spondias philippinensis, Haplospondias other genera [4]. brandisiana and Spondias bipinnata are uncertain, but they are likely belonging to In the revision of tropical Asian Spondias, Spondias [1]. Airy-Shaw & Forman (1967) [5] lumped Allospondias and Solenocarpus with a rather Min & Barfod (2008) [8] recognized two broadly defined Spondias. In contrast, species of Spondias in China S. pinnata and S. Kostermans (1981, 1991) defined the genera lakonensis, in which S. lakonensis includes of the Spondiadoideae rather narrowly, two varieties, S. lakonensis var. lakonensis maintaining Allospondias and Solenocarpus, and S. lakonensis var. hirsuta. transferring Spondias philippinensis (Elmer) Chayamarit (1997) [9] studied phylogeny of Airy-Shaw and Forman to the latter, Anacardiaceae (including Spondias) in describing the new genus Haplospondias and Thailand based on molecular data. However, formally returning the South Pacific species taxon sampling and sequences (only rbcL) of Spondias dulcis Parkinson into the pre- this study were limited. The result of the existing genus Evia Comm. ex Blume emend. study showed close relationship between Kosterm [6], [7]. Spondias and Dracontomelon. Michell & Daly (2015) [1] suggested that Silva et al., (2015) [2] conducted a Allospondias lakonensis (Pierre) Stapf (syn.: phylogenetic study for neotropical species of Spondias lakonensis Pierre var. lakonensis) the genus Spondias. Six species of Spondias should be removed from Spondias based on from neotropic were sampled and three the structure of leaves and flowers such as: makers rbcL, matK and trnH-psbA spacer lack of an intramarginal vein and presence of were applied. The result indicated that perpendicular epimedial tertiary veins, styles neotropical Spondias were divided into two connivent at anthesis and stigmas extrorse on clades. The first clade includes, Spondias the developing fruit, lack of a fibrous matrix mombin and S. purpurea L. while the second on the endocarp. The morphology of S. clade includes, S. cytherea Sonn., S. tuberosa 34 http://jst.tnu.edu.vn; Email: [email protected] Lê Chí Toàn và Đtg Tạp chí KHOA HỌC & CÔNG NGHỆ ĐHTN 207(14): 33 - 40 Arruda, S. venulosa (Engl.) Engl. and [15]. The sequences were aligned in Geneious Spondias sp. However, S. cytherea was v.8.0.5 [15]. distributed widely in the world, thus the 2.2. Morphological analyses neotropical Spondias is likely not a The specimens or photos of specimens of monophyletic. Spondias from the following herbaria: HN, Nguyen (2004) [10] suggested that HNU, PE, HAL, TCD, L, C, A and KUN Vietnamese Spondias includes three species were examined. The herbarium code follow S. cytherea, S. petelotii, S. pinnata and is the Index Herbariorum mainly distributed in some provinces of (http://sweetgum.nybg.org/ih/). We also northern and southern Vietnam such as: Lang observed specimens from herbaria of Son, Lai Chau, Son La, Hoa Binh, Lam Dong deparment of Botany – Ha Noi Pedagogical and Dong Nai. Members of Vietnamese University No. 2 (*) and National Institute of Spondias have significantly economic uses Medicinal Materials (**). Additionally, we and are widely planted. However, Pham examined living materials in the field. (2003) [11] suggested that Vietnamese 2.3. Phylogenetic analyses Spondias includes three species S. pinnata, S. Both maximum likelihood (ML) and cythera and S. mombin. Moreover, the author Bayesian inference (BI) methods were also noted that he did not observed S. mombin employed for the phylogenetic analyses of in Vietnam. Up to now, the studies of Vietnamese Spondias. The ML trees were taxonomy and phylogeny of Vietnamese generated by performing a rapid bootstrap Spondias are limited, thus the relationship analysis in RAxML v.8.2.8 [16], [17] with the between Spondia species in Vietnam is still GTR + I + G substitution model applying unclear and merits further morphological and 1000 bootstrap replicates. The best-fitting molecular analyses. models for the combined datasets were The present study aims to: (1) infer the determined by the Akaike information phylogenetic relationships within Vietnamese Criterion (AIC) as implemented in Spondias, (2) investigate the morphology and jModelTest v.2.1.6 [18]. The Bayesian provide a phylogenetically based analysis was performed in MrBayes v.3.1.2 classification and integrating evidence from [19] on the CIPRES Science Gateway Portal both molecular and morphological data. [20] based on the same models as in the ML 2. Material and methods analysis. The Markov chain Monte Carlo (MCMC) algorithm was run for 5,000,000 2.1. Sampling, DNA extraction, generations with a total of four chains, amplification and sequencing starting from a random tree and trees were The present study sampled 14 species (16 sampled every 1000 generations.
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