MINISTRY OF EDUCATION VIETNAM ACADEMY OF AND TRAINING SCIENCE AND TECHNOLOGY

GRADUATE UNIVERSITY OF SCIENCE AND TECHNOLOGY

------

Nguyen Xuan Quyen

STUDY ON OF FAMILY ( R. Br.) IN VIETNAM

Major: Botany Code: 9.42.01.11

SUMMARY OF BIOLOGY DOCTORAL THESIS

Hanoi, 2021

The thesis is completed at: GRADUATE UNIVERSITY OF SCIENCE AND TECHNOLOGY, VIETNAM ACADEMY OF SCIENCE AND TECHNOLOGY

Supervisors: 1. Tran Thi Phuong Anh Ph.D. 2. Nguyen The Cuong Ph.D.

Examination board Commenter 1: Commenter 2: Commenter 3:

This doctoral thesis will be defended at the GUST-level Board of Examiner at Graduate University of Science and Technology, Vietnam Academy of Science and Technology at ...... on ...... /...... /…….

This doctoral thesis can be found at: - National Library of Vietnam - Library of Gradute Univesity of Science and Technology

INTRODUCTION 1. Rationale for the study Vietnam is located in the tropical monsoon climate, geographic location with complex terrain and many different ecological regions, so the flora is very diverse and rich. From the late 18th century up to now, there have been many studies on taxonomy in our country, including new research results that have contributed to the necessary scientific basis for a number of related fields such as: conservation of biodiversity, ecology, agriculture, forestry, medicine, ... and management such as planning, building economic development, ... In the world, there are many completed and systematic work on plant taxonomy, which are the National Floras. That is the most up-to-date document on the specimen and latest information, using the modern methods of classification in each country, ... In Vietnam, 21 volumes The Flora of Vietnam have been published (2000- 2017), which included 3639 species, 665 genera belonging to 57 families of plant. These are basic scientific achievements of great significance and value in socio- economic development for the country. In the flora of Vietnam, although the Cashew family (Anacardiaceae) do not comprise many species. But it comprises a lot of species with high economic and valuable in use (timber tree, eatable fruits, medicine, latex for paint, oil and fat,...). Up to now, there have been a number of works on taxonomy of the Cashew family, however there are certain limitations, due to not timely updating of information, lack of specimens, incomplete and system. Therefore, the thesis topic "Study on taxonomy of Cashew family (Anacardiaceae R. Br.) in Vietnam" is a work with content meeting the urgent requirements of compiling the Cashew family in the Flora of Vietnam that contribute to the current socio-economic development of the country. 2. The purpose of the thesis topic Completely and systematic study of taxonomy of the Cashew family (Anacardiaceae) in Vietnam, based on evidence of morphology and molecular biology, as a scientific basis for the compilation of Flora of Vietnam about Anacardiaceae in Vietnam. 3. Scientific and practical significance of the thesis topic Scientific significance: Research results supplement and complete the scientific research on taxonomic of the Cashew family (Anacardiaceae) in Vietnam, contributing to the compilation of Flora of Vietnam about Anacardiaceae. Practical significance: Research results serve as a scientific basis for some of related subjects such as forestry, pharmacology, plant resources, biodiversity, ... and training purposes. 4. Structure of the thesis The thesis consists of 150 pages, 9 tables, 80 pictures, 128 photos (123 species), 32 maps. 1

The thesis consists of parts: Introduction (2 pages); Chapter 1. Literature review (29 pages); Chapter 2. Objects, content and research method (8 pages); Chapter 3. Research results and discussion (104 pages); Conclusion, recommendations, and new points of the thesis (2 pages); List of tables (9 tables); List of photos (123 photos); List of pictures (80 pictures); List of maps (32 maps); List of abbreviations (1 page); List of abbreviations of herbaria (2 pages); List of thesis related publications (1 paper); References (8 papers); Lookup table scientific name; Lookup table in Vietnamese name; Appendix 1. Color photos of morphological characteristics of Anacardiaceae in Vietnam (45 photos); Appendix 2. Standard sample photo of Anacardiaceae in Vietnam (39 photos); Appendix 3: Sample image of research of Anacardiaceae in Vietnam (39 photos); Appendix 4: Map of distribution of species of Anacardiaceae family in Vietnam (32 maps); Appendix 5: List of Anacardiaceae family species in Vietnam for genetic sequencing and gene sequencing data (ITS) (17 pages). CHAPTER 1. LITERATURE REVIEW 1.1. The taxonomic position of the Anacardiaceae in the Magnoliophyta classification system Since its establishment (1818) by R. Brown, Anacardiaceae has been changed its position in the classification systems. At first, some authors considered Anacardiaceae to be independent or including Burseraceae. However, the view of Anacardiaceae as an independent family has been confirmed and placed in Magnoliopsida, belonging Magnoliophyta. Some authors classify Anacardicaceae belonging to Rutales, but most classify this family into . 1.2. The systems of Anacardiaceae in the world and the surrounding areas with Vietnam 1.2.1. The ystems of Anacardiaceae in the world The authors divided Anacardiaceae directly to genera: Linnaeus (1753) was the first author to classify and describe 14 species belonging to three genera , Rhus and Anacardium. P. Browne (1756) added Comocladia; genus Astronium, genus were supplemented by J. Jacquin (1760) and Linneaus (1771) and many other authors added more genera. The authors divided the Anacardiaceae directly to tribus: G. Bentham & J. D. Hooker (1887) based on the number of locular of ovary dividing the Anacaediaceae into 2 tribus and 46 genera: tribute Anacardieae (1 locular) comprises 35 genera; and tribute Spondieae (2-5 locular) comprises 11 genera. A. Engler (1892) based on the morphology of flowers, fruits, stamens and ovules divided into 5 tribus: Anacardieae, Spondieae, Rhoideae, Semecarpeae and Dobineeae. A. Engler (1896, 1903) based on anatomical form, detailed description, complete drawings, dividing this family into 5 tribus, 58 genera: Trib. 1. Mangiferae (ovary 5 separate locular), Trib. 2. Spondieae: (ovary 4-5 locular), Trib. 3. Rhoideae (ovary 3 separate locular), Trib. 4. Semecarpeae (ovary 3 clocular) and Trib. 5. Dobineeae (ovary 1 locular). 2

Melchior (1964) based on the Engler’s System dividing Anacardiaceae into 5 tribus: Anacardieae, Spondieae, Rhoideae, Semecarpeae and Dobineae. V. H. Heywood (1996) divided Anacardiaceae into 4 tribus: Trib. 1. Anacardieae (ovary 5 locular, detached, leaf pinnately compound or 1 locular, leaves simple) has 8 genera; Trib. 2. Spondieae (ovary 4-5 locular) has 21 genera; Trib. 3. Semecarpeae (ovary inferior, 3 locular) has 6 genera and Trib. 4. Rhoideae (ovary superior, 3 locular) has 42 genera. So there are 3 authors divided into 4-5 Tribus, only 1 author divided into 2 Tribus. The authors divide Anacardiaceae family into subfamily and tribus A. Takhtajan (1996) divided Anacardiaceae into 4 subfamilies: Subfam. 1. Anacardioideae: (ovary with 5 locular or 1 locular, simple leaf); Subfam. 2. Spondioideae (ovary with 3-5 locular or 1 locular, pinnately compound leaf); Subfam. 3. Juliannioideae (ovary with 1 locular, flower reduced). A. Takhtajan (2009) recorded Anacardiaceae with 78 genera, and about 900 species, which was divided into 4 subfamilies: Subfam. 1. Anacardioideae (5 free carpels or 1 carpel, simple leaf); Subfam. 2. Spondioideae (3-5 united carpels, pinnately compound leaf) with 3 tribus (Spondieae, Rhoeae and Semecarpeae); Subfam. 3. Juliannioideae (ovary with 1 locular, 1 ovule, flower reduced); Subfam. 4. Pistacioideae (3 united carpels, 2 carpels reduced, ovary 1 locular, 12 ovules). S. K. Pell & al. (2011) divided Anacardiaceae in to two subfamilies: Subfam. 1. Anacardioideae (fruits sometimes with wings, ovary 1-3-5 locular) comprises 61 genera; Subfam. 2. Spondioideae (fruits with wings, ovary 4-5 locular) comprises 20 genera. There are also some of other authors divided into subfamilies according to the above point of view, such as Gundersen (1950), Cronquist (1981), Wannan (1986), Thorne (1992), Pell & al. (2001, 2004) ... 1.2.2. Study on classification of Anacardiaceae in surrounding areas with Vietnam Ding Hou (1978) divided the family Anacardiaceae in Malesia into 4 tribus, 22 genera; T. L. Ming (1980) divided Anacardiaceae in China into 5 tribus and 17 genera; J. D. Hooker & C. B. Clarke (1876, 1879) dividesd Anacardiaceae in India into 2 tribus, 21 genera; H. Lecomte (1908) divided Anacardiaceae in Indochina into 9 genera, 13 species; Tardieu - Blot (1962) divided Anacardiaceae in Cambodia, Laos and Vietnam into 12 genera, 18 species; Hui-Lin Li (1993) divided Anacardiaceae in Taiwan into 4 genera, 9 species; A. Backer and R. C. Bakhuizen (1965) divied Anacardiaceae in Java (Indonesia) into 13 genera, 26 species; A. S. George and D. J. Du Puy (1993) studied in Australia with 3 genera with 3 species; F. Likuo & H. Tao (2001) studied in China with 16 genera, 43 species; in Yunnan (China) according to one author (1991) there are 13 genera with 31 species; K. Chayanarit (2010) divided Anacardiaceae in Thailand into 19 genera, 65 species;…

3

Study on classification of Anacardiaceae in Vietnam J. Loureiro (1790) was the first author who study on taxonomy of Anacardiaceae in the Southern Vietnam. He recorded 4 genera, 5 species belonging to Anacardiaceae. Based on the character of stamens and stigma, he divided these species into 2 classes and 2 groups (group with 5 stamens, 1 pistil has 2 genera, 3 species and group with 5 stamens, 3 pistils have 2 genera, 2 species ); L. Pierre (1897, 1898) studied in the Southern Vietnam, he recorded 11 genera and 40 species belonging to Anacardiaceae. H. Lecomte (1908) recorded 14 genera with 31 species and 2 varieties belonging Anacardiaceae in Vietnam. Tardieu-Blot (1962) divided Anacardiaceae in Vietnam into 4 tribus, 19 genera, 57 species and 4 varieties. Vietnamese authors have also had some works on the Anacardiaceae family, but mainly based on the above documents on classification system and basic information, with additional updates on new distribution, use value, ... That is L. K. Ke et al. (1971), many authors at the Forest Inventory and Planning Institute (1971-1988), P. H. Ho (1972, 2000) described and drawing for 70 species, N. T. Ban (2003),... There are also some of authors studied on use of species belonging to Anacardiaceae such as: T. C. Khanh (1992), D. H. Bich et al. (2004), D. T. Loi (2000), V. V. Chi (1991, 1997, 2012), N. T. Ban et al. (2000, 2007), Institute of Medicinal Materials (2016),... 1.3. Situation of research and classifying plants by molecular biology method Recently, plant classification majors still use the comparative morphological method which is basic to determine the scientific names of taxa, but has combined molecular biology methods to support, increase the accuracy of the familiar relationship of taxa. So far, there have been many studies on Anacardiaceae using additional molecular biology methods like Wannan & al. (2006), Pell & Urbatsch (2001), Jessica Silva (2015), ... A number of authors have built plants of phylogenetic trees such as Theodor & al. (2019). In Vietnam, there have been some plant classification works using molecular biology methods such as Vu Cong Hau (2000), Quang Ngoc Vang (2005), Huynh Truong Hue (2008), Vu Thi Thu Hien - Dinh Thi Phong et al. (2012),… 1.4. Commenting on the research situation of the Anacardiaceae family - Summary of studies on the position of the Anacardiaceae family in Magnoliophyta, the situation of researching the Anacardiaceae family systems in the world with 3 perspectives (system of classification to genus, system of classification through tribus to genus, classification system of subfamily, tribus and genus). - Summarized the situation of researching and classifying the Anacardiaceae family in some areas surrounding Vietnam and in Vietnam.

4

CHAPTER 2. SUBJECTS, CONTENTS AND METHODS OF THE STUDY 2.1. Object, scope, research materials - Research subjects of this thesis were taxa (mainly species) belonging to Anacardiaceae in Vietnam. - The research scope of this thesis was all of the regions in Vietnam. - The research materials were all of the individuals belonging to Anacardiaceae living in the wild, specimens were stored at herbaria (about 1200 specimens belonging about 355 collected numbers), drawings, photos, related publications. - Materials for molecular biology research were samples of species belonging to Anacardiaceae, which collected in Vietnam, including 25 dried leaf samples stored in silicagel. The chosen region genomes were the chloroplast genome: trnL-trnF and rbcL. 2.2. Research content - Researching the literature of Anacardiaceae taxonomy in Vietnam and in the world to determine their position in Magnoliophyta and select appropriate classification system of Anacardiaceae in Vietnam. - Determining the sequence of nucleotides of the studied gene regions; building a phylogenic tree of Anacardiaceae in Vietnam. - Identify typical morphological characteristics of Anacardiaceae in Vietnam. - Building the keys to genera and species belonging to Anacardiaceae in Vietnam. - Presentation of information of tribus, genera and species including: Tribus includes scientific name, Vietnamese name, recognizable features, Typus, number of genera and species; The genus includes scientific names, citations, Vietnamese names, synonym, identifiable features, typus, number of species, key to species identification (if any); for species including scientific names, citations, synonym, Vietnamese names, identifiable features, the first sample collection site and types, phenology and habitat, distribution, research samples, validity (if any), notes and comments (if any); accompanied by drawings, photos, distribution maps; comments on validity and distribution of the Cashew family. 2.3. Research methods 2.3.1. Inheritance method and document review: study on related works, specimen in herbarium and research results of Cashew family classification system in the world and Vietnam. 2.3.2. Methods of survey and collection samples in the field: Investigate and collect specimens and documents throughout the country, paying attention to where conditions are right for the Cashew family. 2.3.3. Method of determining scientific names, evaluating scientific values (conservation) and using, building distribution maps, building taxon identification keys: This is a traditional and popular method that are accurate and scienctific. Using

5 monograph classification documents of Cashew family in the world and Vietnam. The morphological term is unified. 2.3.4. Method of molecular biology: Research samples were analyzed at the Department of Molecular Systems and Conservation Genetics, Institute of Ecology and Biological Resources. Including content of total DNA extraction, primer design for sequencing reading, PCR on gene amplification and agarose gel electrophoresis, reading genetic sequence, building phylogenetic tree. CHAPTER 3. RESULTS AND DISCUSSION 3.1. Morphological characteristics of the Anacardiaceae family in Vietnam 3.1.1. Stem: Trees, shrubs and woody climbers, evergreen, resiniferous. 3.1.2. Leaves: Simple or compound leaf; imparipinnate or paripinnate compound leaf; simple leaf and leaflets opposite or alternate or nearly clustered terminal. Leaf blade elliptic, obovate, lanceolate, oblanceolate or oblong; size change; apex acute to acuminate, rounded, obtuse, base obtuse or rounded. 3.1.3. Inflorescences: Terminal or axillary, panicles or raceme. 3.1.4. Flowers: Bisexual or unisexual, 5–merous. 3.1.5. Fruits and seeds: Drupaceous, sometimes borne on enlarged fleshy hypocarp formed by pedicel and receptacle; sometimes winged; pericarp with 3 layers; seed elliptic, ovate, reniform. 3.2. The result of decoding the genetic sequence data analyzed to build a diagram of possible relationship between the genera of Anacardiaceae family in Vietnam 3.2.1. Research samples and information on Genbank: The total number of samples used is 25 dried leaf samples of 15 species that have been scientifically identified. 3.2.2. The sequencing results of gene regions: TrnL-trnF gene region consists of about 1000 pairs of nucleotides; rbcL consists of approximately 600 nucleotide pairs. 3.2.3. Phylogenic tree From the sequencing results, the diagram of the phylogenetic tree genome trnL- trnF and rbcL Anacardiaceae family in Vietnam was built according to ML method, which clearly shows taxonomic units belonging to the genus of Anacardiaceae family evolve according to a separate evolutionary branch, not the same evolutionary branch. Diagram of a phylogenetic tree divided into 4 branches including 4 different branches of the Anacardiaceae family : Branch 1 (Tribus 1) including the genera Mangifera, Gluta, Ancardium, and Bouea are closely related; Branch 2 (Tribus 2) includes the genera Allospondias, Spondias, Choerospondia, Dracontomelon and Pegia belonging to the same branch and are closely related; Branch 3 (Tribus 3), two genera Rhus and Toxicodendron are closely related; Branch 4 (Tribus 4) includes the genera Semecarpus and Drimycarpus are closely related (figures 3.14, 3.15).

6

Figure 3.14: Diagram of a phylogenetic tree (Anacardiaceae), based on the gene rbcL by the Mximum Likelihood method.

Figure 3.15: Diagram of a phylogenetic tree (Anacardiaceae), based on the gene trnL-trnF by the Maximum Likelihood method. 7

3.2.4. Some comments on the result of the above sequence data The sequence of nucleotides of the chloroplast genome trnL-trnF and rbcL shows the genetic differences of taxon as well as research samples of the same morphological species but different distribution areas. The phylogenetic tree diagram shows that the genus Buchanania and the genera Mangifera, Bouea, Gluta of Anacardieae in both trnL-trnF and rbcL genomes are closely related. Thus, molecular biological evidence indicates that the limb and the relation of the genera in the Anacardieae are completely consistent with the trib. Classification system of the authors Engler (1892, 1896, 1903), Menchior (1964), Heywood (1996). The genera Allospondias, Dracontomelon, Spondias, Lannea and Choerospondias are in the same branch and closely related. Combining the two branches, the genera Allospondias and Spondia are more closely related to each other. This is consistent with evidence-based systems of the morphology of these genera of Spondieae. The genera Rhus, Pistacia and Toxicodendron are closely related. The genus Schinus is located between the two genera Rhus and Toxicodendron, the taxa of the genus Rhus are closely related to the genus Schinus. The extra branch of Toxicodendron shows that other samples of the same morphology of Toxicodendron succedaneum that are genetically different when they differ in distribution. The limits and relationship of these genera according to molecular biology evidence are consistent with the evidence-based system of these two genera belonging to Rhoideae. The Semecarpus genus has the same evolutionary branch and is closely related to the taxonomic units of the genus Drimycarpus. Regarding the limit and the relationship of these two genera according to molecular biological evidence consistent with the morphological evidence system of these two genera belonging to Semecarpeae. Phylogenetic tree Anacardiaceae family in Vietnam is suitable for a classification system for Anacardiaceae family. 3.3. Selecting the classification system of Anacardiaceae in Vietnam After analyzing and comparing the systems of Anacardiaceae in the world, based on the combination of morphological and molecular biology data, we selected the classification system of Anacardiaceae in Vietnam based on the system of Engler (1892, 1896, 1903), Menchior (1964), Heywood (1996), to arrange taxonomic units including 4 tribus, 22 genera, 66 species, 6 varieties (table 3.3). Table 3.3. Anacardiaceae family taxonomy system in Vietnam (according to Menchior (1964); Engler (1892, 1896, 1903), Heywood (1996) (4 tribus, 22 genera, 66 species, 6 varieties)

8

Tribus (Trib.) Genus (Gen.) Trib. 1. Anacardieae 1. Melanorrhoea (2 species, 1 var.) (7 genus, 32 species, 5 var.) 2. Buchanania (5 species) 3. Anacardium (1 species) 4. Gluta (6 species) 5. Bouea (2 species, 2 var.) 6. (4 species) 7. Mangifera (12 species, 2 var.) Trib. 2. Spondieae 8. Pegia (1 species) (7 genus, 12 species) 9. Lannea (1 species) 10. Dracontomelon (5 species) 11. Choerospondias (1 species) 12. Pleiogynium (1 species) 13. Spondias (2 species) 14. Allospondias (1 species) Trib. 3. Rhoideae 15. Pentaspadon (2 species) (5 genus, 9 species, 1 var.) 16. Sgenusnus (1 species) 17. Rhus (1 species, 1 var.) 18. Toxicodendron (3 species) 19. Pistacia (2 species) Trib. 4. Semecarpeae 20. Semecarpus (11 species) (3 genus, 13 species) 21. Drimycarpus (1 species) 22. Holigarna (1 species) 3.4. Keys and descriptions of taxa belonging to Annacardiaceae in Vietnam ANACARDIACEAE R. Br. – CASHEW FAMILY R. Br. 1818. Narr. Exped. Zaire: 431; Engl. in De Candolle, 1883. Monogr. Phan. 4: 172; Bark (1942). Am. Mid. Nat. 28: 465; Lecomte (1908), Fl. Gén. Indoch. 2: 6; Tard.-Blot, 1962. Fl. Cambod. Laos Vietn. 2: 67; Phamh. 2003. Illustr. Fl. Vietn. 2: 363; N. T. Ban (2003). Checkl. Pl. Spec. Vietn. 2: 941; T. L. Ming & Anders Basfor, 2008. Fl. China, 11: 335; K. Chayamarit, 2010. Fl. Thailand, 10(3): 265; S. K. Pell & al. in Kubitzki, 2011. Fam. Gen. Vasc. Pl. 10: 7. Typus: Anacardium L. About 81 genera and 800 species, mainly tropical jungle in two hemispheres, sometimes temperate in the world; heliophytes. There are 4 tribus, 22 genera, 66 species and 6 variates in Vietnam. Key to tribus and genera 1A. Ovary with free carpels, sometimes 1-carpel developed .....Trib. 1. Anacardieae 2A. Stamens 30-35, arranged in many whorls …………1. Melanorrhoea 2B. Stamens equal to or petals in number, arranged in 1-2 whorls. 3A. Stamens twice petals in number, (8-10), arranged in 2 whorls 9

4A. Flowers bisexual, stamens equal; without staminode, ovary enclosed by disk; peduncle caduceus ..………………….…………………………..2. Buchanania 4B. Flowers bisexual or unisexual; female flower or bisexual with staminode, 1 stamen fertility; without disk; peduncle fleshy, accrescent cover a portion of the fruit…………..…………………………………………………..3. Anacardium 3B. Number of stamens equal to petals (4-5), arranged in 1 whorl 5A. Flowers bisexual, without staminode 6A. Leaves alternate; flower without disk………………...…………...4. Gluta 6B. Leaves opposite or verticillate; ovary enclosed by disk ...... 5. Bouea 5B. Flowers bisexual or unisexual; female flower or bisexual with staminode 7A Flowers with disc outside stamen; petal accrescent with fruit into wings; endocarp without crustaceous…………………………………….6. Swintonia 7B. Flowers with ovary enclosed by disc; petal not accrescent with fruit; endocarp crustaceous……………………...……….………………....7. Mangifera 1B. Ovary with connated carpels 8A. Ovary with 4-5 connated carpels connate, 4-5 locular. ……...Trib. 2. Spondieae 9A. Lianas………………………………………………………………...8. Pegia 9B. Trees 10A. Perianth 4- merous……………………………….……………...9. Lannea 10B. Perianth 5- merous 11A. Perianth imbricate 12A. Flowers bisexual……………………...………...... 10. Dracontomelum 12B. Flowers unisexual (polygamo-dioecious or dioecious) 13A. Female flower auxillary, solitary or 2-3 flowers; male flower panicles……………………………………………….11. Choerospondias 13B. Female and male flower panicles …………...... 12. Pleiogynium 11B. Perianth valvate 14A. Endocarp crustaceous; lateral veins gradually extending create intramarginal veins...…………………………………………...13. Spondias 14B. Endocarp not crustaceous; lateral veins do not create intramarginal veins ……………………………………………………….....…14. Allospondias 8B. Ovary with 1-3 connated carpels connate, 1 locular. 15A. Leaf pinnate compound, superior ovary ..……………...….Trib. 3. Rhoideae 16A. Perianth with petal 1 whorl, sepal 1 whorl 17A. Number of stamens twice the number of petals (8-10), 2 whorls 18A. Flowers bisexual; drupaceous, oblong ovoid ………15. Pentaspadon. 18B. Flowers unisexual; drupaceous globose…………………….16. Schinus 17B. Number of stamens equal petals (4-5), 1 whorl. 19A. Inflorescences terminal; exocarp ciliate and glandular hair ………………………………………………………………….…..17. Rhus 10

19B. Inflorescences auxillary; exocarp smooth…...... 18. Toxicodendron 16B. Perianth completely reduced or left calyx 1 whorl ...……….…19. Pistacia 15B. Half-inferior or inferior ovary. Peduncle accrescent. Leaves simple ……...……………………………………………………...Trib. 4. Semecarpeae 20A. Half-inferior ovary………………...………………..……..20. Semecarpus 20B. Inferior ovary; 21A. Flowers polygamous; perianthimbricate; stigma 1 ..…21. Drimycarpus 21B. Flowers unisexual; perianth valvate; stigma 3 (female flower) ……………………………………………………...... 22. Holigarna Trib. 1. ANACARDIEAE Typus: Annacardium L. Gen. 1. MELANORRHOEA Wall. Wall. 1829. Pl. Asiat. Rar. 1: t. 11-12. Typus: Melanorrhoea usitata Wall. Key to species 1A. Young branches, leaf abaxial, inflorescences pubescent; fruit drupaceous winged accrescent, 2 times longer than the fruit ……………………………….….1. M. usitata 1B. Young branches, leaf abaxial, inflorences glabrous; fruit drupaceous winged accrescent equal to or shorter than the fruit …...…...………………....…2. M. laccifera 1.1. Melanorrhoea usitata Wall. Wall. 1829. Pl. Asiat. Rar. 1: 9. Loc.class.: India. Typus: Wallich CAT 597 (BR: isotype, BR 0000006954291, photo!) 1.2. Melanorrhoea laccifera Pierre Pierre, 1897. Fl. For. Cochinch. tab. 367A. Loc. class.: Cambodia (Somrong Tong). Typus: Pierre 915 (P: isotype, MPU 20608; photo!) 1.2.a. Melanorrhoea laccifera var. parvifolia Evrard Evrard, 1952. Bull. Soc. Bot. France, 99: 131. Loc. class.: Vietnam, Khanh Hoa. Typus: Poilane 4573 (P). Gen. 2. BUCHANANIA Spreng. Spreng, in Schrad. 1818. Journ. Bot. 2: 234 Typus: Buchanania lanzan Spreng. Key to species 1A. Anthers lanceolate ………………………………………….…..1. B. arborescens 1B. Anthers are not lanceolate 2A. Leaves hairy………………………...…………………….2. B. cochinchinensis 2B. Leaves glabrous (or sparse hair in abaxial) 3A. Secondary veins densely pinnate, conspicuous adaxial; flower sessile or short; inflorescences raceme …………………………………………....3. B. reticulata

11

3B. Secondary veins not pinnate in adaxial or inconspicuous; peduncle equal to bud; inflorescences panicles 4A. Leaves large, 10-25 cm at length; inflorescences shorter than leaf, outer sepal glabrous…………...……………………………………..……..4. B. glabra 4B. Leaves small, 4-9 cm at length; inflorescences equal to or longer than leaf, outer sepal hairy……...………………………………….……...5. B. siamensis 2.1. Buchanania arborescens (Blume) Blume Blume, 1850. Mus. Bot. Lugd.-Bat. 1: 183 Loc.class.: Indonesia (Java). Typus: C. L. Blume sine num. (L, L 001552, photo!). 2.2. Buchanania cochinchinensis (Lour.) M. R. Almeida M. R. Almeida, 1996. Fl. Maharashtra, 1: 287. Loc.class.: Indonesia. Typus: Roxburgh 103 (K) 2.3. Buchanania reticulata Hance Hance, 1877. Journ. Bot. 15: 332; Loc.class.: Philippines (Sibuyan). Typus: ADE Elmer 12334 (L: isotype, L 001567, photo!). 2.4. Buchanania glabra Wall. ex Engl. Engl. 1883. Monogr. Phan. 4: 183 Loc.class.: India (Moulmein). Typus: N. Wallich 984 (A, A 00049012, photo!). 2. 5. Buchanania siamensis Miq. Miq. 1869. Ann. Mus. Bot. Lugduno-Batavi, 4: 118 Loc.class.: Thailand (Radboerei). Typus: Teysman sine num. (L, L 005583, photo!). Gen. 3. ANACARDIUM L. L. 1753. Sp. Pl.l: 383 Typus: Anacardium occidentale L. The world has about 10 species. In Vietnam, one species is recorded. 3.1. Anacardium occidentale L. L. 1753. Sp. Pl. 1: 383 Loc. class.: Sri Lanka. Typus: P. Hermann sine num. (BM, lectotype, BM 000621986, photo!). Gen. 4. GLUTA L. L. 1771. Mant. Pl. 2: 293 Typus: Gluta renghas L. Key to species 1A. Inflorescences and flower hairy 2A. Fruit drupaceous big, 3-5 cm in dia. 2B. Fruit drupaceous small, 0,3-0,5 mm in dia…………………...…… 1. G. gracilis 3A. Peel rough; aerial roots………………………………………..…..2. G. velutina 3B. Peel smooth, no aerial roots 4A. Ovary hairy…………………………………………………….…3. G. wrayi 12

4B. Ovary glabrous 5A. Fruit drupaceous ovoid or reniform …………………...4. G. megalocarpa 5B. Fruit drupaceous, globose …………….…………………….5. G. tavoyana 1B. Inflorescences and flower glabrous………………………….…….6. G. compacta 4.1. Gluta gracilis Evrard Evrard, 1952. Bull. Soc. Bot. France, 99: 83 Loc. class.: Vietnam (Cana province de Phanrang). Typus: Poilane 9004 (P: holotype, P 02440674, photo!; Isotype, HM! & P, P 02440675, Photo!). 4.2. Gluta velutina Blume Blume, 1850. Mus. Bot. 1: 183 Loc. class.: Indonesia (Sumatra). Typus: Mullar sine num. (L, L 0015644, photo!). 4.3. Gluta wrayi King King, 1896. Journ. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 65: 482 Loc. class.: India, Perak. Typus: Wray 2290. 4.4. Gluta megalocarpa (Evrard) Tardieu Tardieu, 1961. Adansonia s.n. 1: 195 Loc. class.: Vietnam, Nha Trang (Hon Heo mountain). Typus: Poilane 4796 (P). 4.5. Gluta tavoyana Wall. ex Hook. f. Hook. f. 1876. Fl. Brit. India, 2: 22 Loc. class.: Malaysia (Tavoy). Typus: Wallich CAT 1004. (BR, Isotype, BR 0000006952884, photo!). 4.6. Gluta compacta Evrard Evrard, 1952. Bull. Soc. Bot. France, 99: 84 Loc. class.: Vietnam, Khanh Hoa (Masif de Co Inh près de Nha Trang). Typus: Poilane 4630 (P: Holotype, P 02440671 photo!; Isotype, P 02440672 & P 02440673, photo!). Gen. 5. BOUEA Meissn. Meissn. 1837. Pl. Vasc. Gen. 1: 75 Typus: Bouea oppositifolia (Roxb.) Meissn. The world has about 5 species. In Vietnam recorded 02 species and 2 var. Key to species 1A. Inflorescences 2 times longer than the leaf; petioles to swell, with cork cambium, pinnate veins conspicuous both surface ……………………………..…..1. B. poilanei 1B. Inflorescences shorter than leaf; petiole flat, no cork cambium; pinnate veins inconspicuous both surface ……………………………………..…..2. B. oppositifolia 5.1. Bouea poilanei Evrard Evrard, 1952. Bull. Soc. Bot. France, 99: 132 Loc. class.: Vietnam, Cochinchine (Dong Nai, Bien Hoa: Trang Bom). Typus: Poilane 23594 (Syntype – L, L 0015541 photo!; K, K 000695211 photo!). 5. 2. Bouea oppositifolia (Roxb.) Meissn. Meissn. 1837. Pl. Vasc. Gen. 2: 55 13

Loc. class.: Myanmar. Typus: Wallich 5.2a. Bouea oppositifolia var. roxburgii (Pierre) Tard.-Blot. Tard.-Blot, 1962. Fl. Cambod. Laos Vietn. 2: 128. Loc. class.: Vietnam, Ba Ria-Vung Tau: Nui Dinh (in montibus Dinh ad Baria). Syntypus: Pierre 123 (HM), Chevalier 36619. (MPU, MPU 020602, photo!). 5.2b. Bouea oppositifolia var. microphylla (Engl.) Merr. Merr. 1930. Ling. Sci. Journ. 9: 39. Loc. class.: Myanmar. Typus: Wallich (?). Gen. 6. SWINTONIA Griff. Griff. 1846. Proc. Linn. Soc. 1: 283. Typus: Griff . (Swintonia griffithii Kurz) The world has about 16 species. Vietnam recorded 04 species. Key to species 1A. Fruit drupaceous, wing accrescent 2-7 times longer than the fruit 2A. Young branches, leaves, inflorescences densely ferruginous pubescent ……………………………………………….…………….……….....1. S. maingay 2B. Young branches, leaves, inflorescences without ferruginous pubescent. 3A. Leaf 5-7 cm long; petiole is as long as 2/3 of leaf blade ...…...…2. S. minuta 3B. Leaf twice as long as the above species (12-15 cm); petiole is shorter than 1/3 of leaf blade ……………………….………………………………..3. S. griffithii 1B. Fruit drupaceous, winged accrescent shorter 1/4 fruit…..………...……4. S. pierrei 6.1. Swintonia maingay. Phamh. 2003. Illustr. Fl. Vietn. 2. 370 “as Swintonia maingayi”. 6.2. Swintonia minuta Evrard Evrard, 1952. Bull. Soc. Bot. France, 99: 84. Loc. class.:Vietnam, Khanh Hoa (?). Typus: Poilane 4957 (P). 6.3. Swintonia griffithii Kurz Kurz, 1870. Journ. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 39: 77 Loc. class.: Mergui. Typus: Helfer 1124 (C, C 10005531, photo!). 6.4. Swintonia pierrei Hance Hance, 1876. Journ. Bot. 14: 257 Loc. class.: Cambodia (Mounts Cam Chay prope Kamput). Typus: Pierre 1460 (MPU, MPU 020628, photo!). Gen. 7. MANGIFERA L. L. 1753. Sp. Pl. 1: 200 Typus: L. The world has about 69 species. Vietnam recorded 12 species, 2 variates. Key to species 1A. Stamen fertility 5 2A. Inflorescences densely pubescent; petiole 1,5-6 cm, leaf blade 8-25 cm 14

3A. Stamen 4-6 lepidote cause stamen reduced; fruit drupaceous ovoid broad or subglobose, ca. 4-5 × 3-5 cm……………………………....…….1. M. duperreana 3B. Stamen without lepidote cause stamen reduced; fruit drupaceous eliptc, ca. 3 × 1,5 cm………………………………………..….…………..2. M. cochinchinensis 2B. Inflorescences glabrous; petiole 0,4-0,5 cm long, leaf blade 4-5 cm ……………………………………………………………….……3. M. minutifolia 1B Stamen fertility 1-2 4A. Stamen fertility 1 5A Inflorescences and flowers pubescent 6A. Disc reduced, small dots, no hairy; the inner of the petal has a midvein …………………………………...... ……….………………………4. M. caesia 6B. Disc thick, annulus, hairy; the inner of the petals has 3-5 veins 7A. The stigma on the top; seed reniform ………………...5. M. camptosperma 7B. The stigmas deviate to the right side; seed oval or flat ovoid …………………………………………………………....……….6. M. indica 5B. Inflorescences and flowers glabrous 8A. Inflorescences shorter than leaf…………...………………...……7. M. flava 8B. Inflorescences longer than leaf 9A. Disc thick, annulus, enclose ovary, the edge has lobes 10A. Inflorescences above pedicle 1/2 length; petal 6-8 mm …8. M. laurina 10B. Inflorescences nearly sessile; petal shorter than 3 mm………………... …………………………………………………………..9. M. dongnaiensis 9B. Disc almost completely reduced 11A. Leaf apex and base rounded or obtuse, rarely acute apex; lobe sepal ovoid, 2 × 1 mm; petal oval, ca. 4-6 × 1,5 × 2,5 mm ...…..….10. M. odorata 11B. Leaf apex and base acute; lobe sepal ovoid, 4 × 2 mm; petal oblong lanceolate, 7-10 × 1,5-2,5 mm………...………………………11. M. foetida 4B Stamen fertility 2 ………………………….…………....……..……..12. M. reba 7.1. Mangifera duperreana Pierre Pierre, 1897. Fl. Forest. Cochinch. 1: tab. 362 Loc. class.: Vietnam, Phu Quoc. Typus: Pierre 1651 (P: holotype, P 02440613, photo!; isotype, P 02440614- P 02440616, photo!). 7. 2. Mangifera cochinchinensis Engl. Engl. 1883 Monogr. Phan. 4: 205 Loc. class.: Vietnam (Province de Bien Hoa). Syntypus: Thorel 1287 (P: Holotype, P 02440617, photo!. Isotype, P 02440618 & P P 02440619, photo!; HM!). 7.3. Mangifera minutifolia Evrard Evrard, 1952. Bull. Soc. Bot. Fr. 99: 83 Loc. class.: Vietnam, Khanh Hoa (Nui Han, prov. de Nha trang). Typus: Poilane 4803 (P, P 0015742 - P 0015742, photo!; HM!). 15

7.4. Mangifera caesia Jack Jack in Roxb. 1824. Fl. Ind.: 441 Loc. class.: Sumatra. Typus: 7.5. Mangifera camptosperma Pierre Pierre, 1897. Fl. Forest. Cochinch. 1: pl. 363A Loc. class.: Vietnam (Près de Tay Ninh), Typus: Pierre 1683 (P: holotype, P 02440628 photo! & isotype P 02440629, photo!). 7.6. Mangifera indica L. L. 1753. Sp. Pl.: 200 Loc. class.: India. Lectotypus: "Mau" in Rheede, Hort. Malab., 4: 1, t. 1, t. 2 (1683) designated by A. J. Bornstein. Currently our country exists many different types of Mango: Cv. indica L.; Cv. cambodiana Pierre; Cv. mekongensis Pierre; Cv. odorata Griff ; (T. Đ. Lý, 1993; Đ. T. Lợi, 2000; N. T. Bân, 2003; V. V. Chi, 2004, 2012; N. Bá & cs, 2014). 7.6a. Mangifera indica var. mekongensis Pierre Pierre, 1887. Fl. For. Coch. 1: pl. 316A 7.6b. Mangifera indica var. cambodiana Pierre Pierre, 1897. l.c pl. 316B 7.7. Mangifera flava Evrard Evrard, 1952. Bull. Soc. Bot. France,. 99: 82 Loc. class.: Vietnam (Massif du Braian pris de Djiring, Dong nai). Typus: Poilane 24055 (P: Holotype, P 02440623, photo!; isotype, P 02440624 - P 02440626, photo!). 7.8. Mangifera laurina Blume Blume, 1850. Mus. Bot. 1: 195 Loc. class.: Malacca. Typus: Griffith 1096 (K). 7.9. Mangifera dongnaiensis Pierre Loc. class.: Vietnam (Pho Qua, sur les river du Dong Nai, prov. de Bienhoa). Typus: Pierre 1659 (P: holotype, P 02440621, photo! & isotype, P 02440621, photo !). 7.10. Mangifera odorata Griff. Griff., 1854 Not. PI. Asiat. 4: 417 Loc. class.: Malacca. Typus: Griffith s.n. (P, P 02440645 & P 05189926, photo!). 7.11. Mangifera foetida Lour. Lour. 1790. Fl. Cochinch.: 160 Loc. class.: Cochinchin. Typus: Loureiro(?). 7. 12. Mangifera reba Pierre Pierre, 1897. Fl. Forest. Cochinch. 1: tab. 363 B Loc. class.: Vietnam (Dong nai, Biên hoa, Pho qua). Typus: Pierre 1648 (P: holotype, P 02440647, photo!; isotype HM! & P, P 02440648 & P 02440649, photo!). Trib. 2. SPONDIEAE Typus: Spondias L. 16

Vietnam has 7 genus and 12 species Gen. 8. PEGIA Colebr. Colebr. 1827. Transact. Linn. Soc. London, 15: 364 Typus: Pegia nitida Colebr. The world has 2 species. In Vietnam recorded one species. 8.1. Pegia sarmentosa (Lecomte) Hand. - Mazz. Hand.-Mazz. 1933 Sinensia, 3: 187. Loc. class.: Vietnam (Tonkin Son Tay: Tu Phap). Typus: Balansa 3695 (P, lectotype, K 000695353, photo!). Gen. 9. LANNEA A. Rich. A. Rich. in Guillem. 1831. Fl. Seneg. Tent.: 153 Typus: Lannea velutina A. Richard. The world has over 40 species. In Vietnam recorded one species. 9.1. Lannea coromandelica (Houtt.) Merr. Merr. 1938. Journ. Arnold Arbor. 19(4): 353 Loc. class.: Thailand. Typus: A. Richard Gen. 10. DRACONTOMELON Blume Blume, 1850. Mus. Bot. 1: 231. t. 42 Key to species 1A. Calyx pubescent on outer 2A. Bud, 2 mm, calyx 1 mm; ovary pubescent…………………...…1. D. laoticum 2B. Bud 6-7 mm, calyx 4-6 mm; ovary glabrous 3A. Disc long hairs at the edge; sepal triangle, inner glabrous…………2. D. dao 3B. Disc entire or undulate at the edge, glabrous; sepal oval, inner hairy ……………………………………………….……………….3. D. duperreanum 1B. Calyx glabrous. 4A Sepal acute, equal to petal in length.…………………………..…..4. D. schmidii 4B. Sepal rounded, shorter than petal 2-4 times……………….……….... D. petelotii 10.1. Dracontomelon laoticum Evrard & Tardieu Evrard & Tardieu, 1962. Adansonia n.s., 1: 198 Loc. class.: Lao (Phou Bassac). Typus: Poilane 20404 (P). 10.2. Dracontomelon dao (Blanco) Merr. & Rolfe Merr. & Rolfe, 1908. Philipp. Journ. Sci., 3: 108 Loc. class.: Philippine (Taytay). Typus: E. D. Merill 9391 (MO, isotype, MO 57478, photo!). 10.3. Dracontomelon duperreanum Pierre Pierre, 1898. Fl. Forest. Cochinch. 5, pl. 374 B Loc. class.: Vietnam, Dong Nai, Bien Hoa. Typus: Pierre 1864 (P). 10.4. Dracontomelon schmidii Tardieu Tardieu, 1961 Adansonia n.s., 1: 55 (“Dracondonlum”) 17

Loc. class.: Vietnam (Dong Nai: Dinh Quan). Typus: Schmid 1959 (P, P 02440654 & P 02440655 photo!; HM!). 10.5. Dracontomelon petelotii Tardieu Tardieu, 1961. Adansonia n.s., 1: 57 Loc. class.: Vietnam (Lang Son: Dong Mo, Van Linh). Typus: Petelot 6384 (P: holotype; NY: isotype, A 00310939, photo! & HNF!). Gen. 11. CHOEROSPONDIAS B. L. Burtt & A.W. Hill B. L. Burtt & A.W. Hill, 1937. Ann. Bot. (Oxford) n.s. 1: 254 Typus: Choerospondias axillaris B. L. Burtt & A.W. Hill. There is 1 species in the world that distributed in India, Nepal, Myanmar, China, Japan, Thailand, Laos and Vietnam. 11.1. Choerospondias axillaris (Roxb.) B. L. Burtt & A.W. Hill B. L. Burtt & A.W. Hill, 1937. Ann. Bot. (Oxford) n.s. 1: 254 Loc. class.: China (Yunnan). Typus: A. Henry 11690 (K). Gen. 12. PLEIOGYNIUM Engl. Engl. in C. DC. 1883. Monogr. Plan. 4: 255. Typus: Pleiogynium solandri (Benth.) Engl. Vietnam has 1 species 12.1. Pleiogynium timoriense (C. DC.) Leenh. Leenh.1952. Blume, 7: 159 Loc. class.: Timo - Letse. Typus: Lambton sine num. (K, isotype, K 000695633, photo!). Gen. 13. SPONDIAS L. L. 1753. Sp. Pl. 1: 371 Typus: Spondias monbin L. The world has about 10 species. Vietnam recorded two species. Key to species 1A. Flower with pedicel; leaflet margin serrat or crenate……..…………...1. S. dulcis 1B. Flower sessile; leaflet margin entire…………….………..………….2. S. pinnata 13.1. Spondias dulcis Park. Park. 1773. Journ. Voy. South Seas, 39 Loc. class.: India(?). Typus: Roxburgh(?). 13.2. Spondias pinnata (L. f.) Kurz Kurz, 1875. Prelim. Rep. Forest Pegu App. A: 44, App. B: 42 Loc. class.: India. Typus: Konig sine num. (C, C 10005527, photo!). Gen. 14. ALLOSPONDIAS Stapf Stapf, 1900. Hooker's Icon. Pl. 27: t. 2667 Typus: Allospondias lakonensis (Pierre) Stapf The world has 3 species. Vietnam has one species. 14.1. Allospondias lakonensis (Pierre) Stapf Stapf, 1900. Hooker's Icon. Pl. 27: t. 2667. 18

Loc. class.: Siam, Lakoon (Kakone). Typus: Harmand, 1825 (P). Trib. 3. RHOIDEAE Typus: Rhus L. Vietnam has 5 genus, 8 species, and 1 var. Gen. 15. PENTASPADON Hook. f Hook. f. 1860. Trans. Linn. Soc. London 23(1): 168 Typus: Pentaspadon motleyi Hook. f. The world has 6 species. Vietnam has 2 species Key to species 1A. Sepal 20-25 cm; inflorescences 10-15 cm; staminodes reduced to scales; ovary pubescent……………………………………………...…………………1. P. poilane 1B. Sepal 15-17 cm; inflorescences ca. 20 cm; 5 staminodes alternate 5 fertility stamen; ovary glabrous or slightly hairy at the base…………………2. P. annamense 15.1. Pentaspadon poilanei (Evrard & Tardieu) Phamh. Phamh. 1992. Illustr. Fl. Vietn. 2(1): 474 Loc. class.: Vietnam (Annam: Hon Heo près de Nhatrang). Typus: Poilane 6100 (P, photo!; HM!). 15.2. Pentaspadon annamense (Evrard & Tardieu) Phamh. Phamh. 1992. Illustr. Fl. Vietn. 2(1): 474 Loc. class.: Vietnam (Annam: Hon Heo près de Nhatrang). Typus: Poilane 4758 (P, P 02449688, photo!). Gen. 16. SCHINUS L. L. 1753. Sp. Pl.: 388. Lectotypus: L. Vietnam has 1 species 16.1. Schinus terebinthifolius Raddi Raddi, 1820. Mem. Mat. Fis. Soc. Ital. Sci. Modena. Pt. Mem. Fis. 18 (2):399-400 Loc. class.: Braxin. Typus: Raddi sine num. (FI, FI 005127, photo!). Gen. 17. RHUS L. Mull. 1768. Gard. Dict. (ed. 8): 7 Typus: Rhus coriaria L. The world has about 40 species. Vietnam has 1 species and 1 var. 17.1. Rhus chinensis Mill. Mull. 1768. Gard. Dict. (ed. 8): 7 Loc. class.: China. Typus: Chevalier 37887 (P). 17.1a. Rhus chinensis var. roxburghii (DC.) Rehd. & Wils Rehd. & Wils 1914. Journ. Arnold Arbor. 20(4): 416 Loc. class.: China. Typus: Roxburgh Gen. 18. TOXICODENDRON Mill. Mill. 1754. Gard. Dict. Abr. (ed.) 4 19

Typus: Toxicodendron pubescen Mill. Key to species 1A. Branches fat; leaves concentrated at terminal, nearly verticillate, leaf scars conspicuous. Peduncle of fruit fleshy, straight, upwards; thick fruit. 2A. Young branches and abaxial slightly sericeous, white. Fruit drupaceous globose, not flat …………………………………………………………...…1. T. rhetsoides 2B. Young branches and abaxial dense ferruginous pubescent. Fruit drupaceous globose, flattened vertically……………………………………..……2. T. wallichii 1B. Branches slender; leaves alternate, not concentrated at terminal, leaf scars indistinct. Peduncle of fruit slender, hanged down; sparse fruit …………………………………………………….…………...…. 3. T. succedaneum 18.1. Toxicodendron rhetsoides (Craib) Tard.-Blot Tard.-Blot, 1962. Fl. Cambod. Laos Vietn. 2: 188 Loc. class.: Thailand. Typus: Kerr 5169 (Holo. – K, BM 000884623, photo!). 18.2. Toxicodendron wallichii (Hook. f.) Kuntz. Kuntz. 1891. Revis. Gen. Pl. 154 Loc. class.: Temperate Himalaya from Garwhal to Nepal, alt. 6-7000 ft. Typus: Wallich sine num. (C, isotype, C 10005547). 18.3. Toxicodendron succedaneum (L.) Kuntze Kuntz. 1891. Revis. Gen. Pl. 1: 154 Loc. class.: Japonia, China. Gen. 19. PISTACIA L. L. 1753. Sp. Pl. 2: 1108 Typus: Pistacia vera L. Key to species 1A. Leaf imparipinnate compound, leaflet blades 4-8 x 2-3,5 cm …1. P. cucphuongensis 1B. Leaf paripinnate compound, leaflet blades 1-1,5 x 0,8-1 cm …………………….. ………….……………………………………………………….2. P. weinmannifolia 19.1. Pistacia cucphuongensis T. Ð. Ðai T. Ð. Dai, 1981. Journ. Biol. (Vietnam) 3 (3): 24, fig. 2 Loc. class.: Vietnam (Cuc Phuong). Typus: Nguyen Tien Ban 948 (Holo. HN!). 19.2. Pistacia weinmannifolia Poiss. ex Franch. Franch. 1886. Bull. Soc. Bot. France, 33: 467 Loc. class.: China (Sichuan). Typus: Wilson, Veitch Exped. 3367. Trib. 4. SEMECARPEAE Typus: Semecarpus L. f. Vietnam has 3 genera, 13 species. Gen. 20. SEMECARPUS L. f. L. f. 1781. Suppl. Pl. 25: 182 Typus: Semecarpus anacardiopsis L. f. 20

Key to species 1A. Leaves densely pubescent 2A. Inflorescences longer than leaves 3A. Inflorescences densely, 20-40 cm long; peduncle fat, branched, lateral branch at base of inflorescences 20-35 cm long; petiole shorter than 1,5 cm. 4A. Peduncle cover most of the fruit; leaf blade oblong, 20-60 cm, apex acute, base oblong………………………………………...……..…….1. S. tonkinensis 4B. Peduncle cover 1/3 under the fruit; leaf blade obovoid, 10-18 cm long, apex rounded or concave, base lanceolate …………………....…2. S. cochinchinensis 3B. Inflorescence sparse, 20-25 cm long; peduncle slender, few branches, lateral branch at base of inflorescences 2-5 cm; petiole 1,5-2 cm ………………………………………………………………..….3. S. graciliflora 2B. Inflorescences shorter than leaves 5A Petiole 3-4 cm, leaf blade ca. 20-30 × 8-10 cm, apex rounded …………….…………………………………...……………...……..4. S. velutina 5B. Petiole 1 cm, leaf blade ca. 15-20 × 3,5-5 cm, apex caudate- acuminate………………………………………………….…….5. S. annamensis 1B. Leaves glabrous 6A. Inflorescences longer than leaf 7A. Peduncle cover more than 2/3 fruit. ……………..…………..…6. S. reticulata 7B. Peduncle cover less than1/3 8A. Petiole 3-5 cm; peduncle shorter 1/3 fruit, cover 1/4 fruit; fruit drupaceous 1-1,2 x 1,5-2 cm. 8B. Petioles 1-1,5 cm; peduncle nearly equal to fruit, cover 1/3 fruit; fruit drupaceous 1-1,2 x 0,8-1 cm …………………………………..……7. S. humilis 9A. Inflorescences 20-40 cm long, lateral branch at base of inflorescences 15- 20 cm; leaf blade ca.15-25 x 8-12 cm, apex rounded or caudate short …………………………………...……………………..…8. S. anacardiopsis 9B. Inflorescences 15-20 cm, lateral branch short; leaf blade 10-15 x 3,4-4,5, apex acute, caudate...…………………………….…………….9. S. perniciosa 6B. Inflorescences shorter than leaf 10A. Petiole 5-10 cm; calyx and petal tomentose; fruit nearly flat globular, less than 0,5 cm ….…………………………………………………………....…10. S. myriocarpa 10B. Petiole shorter than 1 cm; calyx and petal glabrous; fruit obovoid, 1,5-2 x 1,2-1,6 cm…………………………………………………..…………….....11. S. caudate 20.1. Semecarpus tonkinensis Lecomte Lecomte, 1908. Bull. Soc. Bot. Fr. 54: 609 Loc. class.: Vietnam, Tonkin (Ao Ca). Typus: Bon 2592 (P, HNF!). 20.2. Semecarpus cochinchinensis Engl. Engl. in DC. 1883. Monogr. Phan. 4: 489 21

Loc. class.: Vietnam (Cochinchin Saigon: Cho Lon). Syntypus: Thorel 670 (P, MPU 021086, photo!; HM!), Poilane 8520 (HM!). 20.3. Semecarpus graciliflora Evrard & Tardieu Evrard & Tardieu, 1961. Adansonia. n. sér. 1: 203 Loc. class.: Vietnam (Nha Trang). Typus: Evrard 540 (P). 20.4. Semecarpus velutina King King, 1896. Journ. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 65: 508 Loc. class.: Malaysia (Perak Larut). Syntypus: Poilane 970 (P, HM!), King 7439 (K, K 000695425!), King 7655 (K, K 000695427, photo!). 20.5. Semecarpus annamensis Tardieu Tardieu, 1961. Adansonia n.ser. 1: 203 Loc. class.: Vietnam (Kon Tum). Typus: Poilane 36689 (P, HM!). 20.6. Semecarpus reticulata Lecomte Lecomte, 1907. Bull. Soc. Bot. France,. 54: 610 Loc. class.: Laos (Vientine: MeKong). Typus: Thorel 3262 (P, isotype, MPU 021081, photo!; HM!). 20.7. Semecarpus humilis Evrard & Tardieu Evrard & Tardieu, 1961. Adansonia. n. sér, 1: 204 Loc. class.: Vietnam (Quang Nam: Mang Tro). Typus: Poilane 31721 (P). 20.8. Semecarpus anacardiopsis Evrard & Tardieu Evrard & Tardieu, 1961. Adansonia n.ser., 1: 204 Loc. class.: Vietnam (Quang Tri). Typus: Eberhardt 2779 (P, HM!). 20.9. Semecarpus perniciosa Evrard & Tardieu Evrard & Tardieu, 1961. Adansonia. N. sér. 1: 206 Loc. class.: Vietnam (Ba Na). Typus: Poilane 7081 (P, HM!). 20.10. Semecarpus myriocarpus Evrard & Tardieu Evrard & Tardieu, 1961. Adansonia. n. sér, 1: 204 Loc. class.: Vietnam (Tay Ninh). Typus: Müller 152 (P, HM!). 20.11. Semecarpus caudata Pierre Pierre, 1898. Fl. Forest. Cochinch. t. 369B. Loc. class.: Vietnam (Dong Nai). Typus: Pierre I679 (P, HM!) Gen. 21. DRIMYCARPUS Hook. f Hook. f. in Benth. & Hook. f. 1862. Gen. Pl. 1: 424 Typus: Drimycarpus racemosus Hook. f. Only one species, distributed in the Himalayas, Myanmar, China, Indochina and Vietnam. 21.1. Drimycarpus racemosus (Roxb.) Hook. f. Hook. f. in Benth. & Hook. f. 1862. Gen. Pl. 1: 424 Loc. class.: Nepal, Himalaya(?). Typus: Roxb. 2213 (K). Gen. 22. HOLIGARNA Buch.-Ham. ex Roxb. (nom. cons.) Roxb. 1814. Hort. Beng.: 22 22

Typus: Holigarna longifolia Buchm.-Ham. ex Roxb. The world has about 7-10 species. One species distributed in Indochina and Vietnam 22.1. Holigarna kurzii King King, 1896. Journ. Asiat. Soc. Bengal. Pt. 2. Nat. Hist. 65(2): 512 (“Holigarnia”); Loc. class.: Thailand. Typus: Kurz 2016 (K). 3. 5. Some comments about the Anacardiaceae family in Vietnam - Scientific value: One new recorded species to the flora of Vietnam is Toxicodendron wallichii; 3 threatened species listed in Vietnam Red Data Book (2007); 18 species and 3 variates has been recorded endemic to flora of Vietnam. - Use value: Timber trees: 37 species (51.4%); edible fruit: 28 species and 3 variates (38.9%); medicinal and poisonous plants: 43 species and variates (59.8%%); resins: 10 species (13.9%); for tannins to dye cloth; shade trees, ornamental plants; oily trees; essential oil plants; plants for edible leaves; ... The total number of valuable species used is 62 species, 4 variates (91.7%) belonging to 22 genera. - Distribution characteristics of Anacardiaceae family in Vietnam: species with small distribution points: there are 39 species, 4 var. of 12 genera; species with many distribution points: there are about 7 species belonging to 6 genera; species with distribution range in the North about 9 species belonging to 7 genera; species with distribution range in the South of about 22 species, 4 var. of 12 genera. CONCLUSIONS AND RECOMMENDATIONS 1. The classification system of Engler (1892, 1896, 1903), Menchior (1964) and Heywood (1996) were selected to classify the taxa of the Mango family (Anacardiaceae) in Vietnam, including 66 species, 6 species, belonging to 22 genera, divided into 4 tones (Anacardieae 7 genera, Spondieae 7 genera, Rhoideae 5 genera and Semecarpeae 3 genera). 2. A new species was recorded to the Flora of Vietnam (Toxicodendron wallichii (Hook. F.) Kuntz); identified 18 species, 3 variates belonging to 10 genera are endemic (or sub-endemic) of Vietnam. 3. Keys to genera and species have been built, using the exact opposite characteristics to use for 22 genera, species of 12 genera with 2 or more species of the Cashew family in Vietnam. 4. All species and varieties are studied on a specimen; be revised, supplemented with many updated information (nomenclature, type, morphological characteristics, distribution, phenology, habitat and comments if any; accompanied by drawings, photos, distribution diagrams). 5. The first time, some taxa belonging to Anacardiaceae in Vietnam were sequenced; the phylogeny tree of Anacardiaceae in Vietnam was built. From there, determine the suitability of the traditional classification system (morphological comparison) with the arrangement into 4 tribus or 4 branches.

23

6. Scientific value assessment (conservation): there are 3 species belonging to 2 genera that are rare and endangered, requiring conservation measures; 62 species, 4 variteies belonging to 22 genera are validity; the distribution of species: the North has 8 species of 6 genera, the South has 24 species, 4variteies of 11 genera, popular nationwide there are 7 species of 6 genera, scattered everywhere there are 4 species of 3 genera. Thus, the cashew family is mainly distributed in the South of Vietnam. Recommendations: It is necessary to continue researching and supplementing materials and specimens, combining molecular biology methods to solve a number of doubtful specimens so that there is enough scientific basis to complete the study of taxonomy of Anacardiaceae serving the publication of Flora of Vietnam about this family. NEW POINTS OF THE THESIS - The thesis is the first complete and systematic scientific work on classification of the mango family (Anacardiaceae) in Vietnam, including 66 species, 6 varieties, belonging to 22 genera, classified into 4 tribus (Anacardieae 7 genera, Spondieae 7 genus, Rhoideae 5 genera and Semecarpeae 3 genera); among that One new recorded species for the Flora of Vietnam (Toxicodendron wallichii (Hook. F.) Kuntz) was found as well as identifying some taxon as endemic (or sub-endemic) of our country (18 species, 3 varieties belonging to 10 genera); all species were studied on a specimen basis; at the same time revise and supplement many new information (nomenclature, research samples, morphology, drawings, distribution diagram, photos, phenology, habitat and comments if any,…). - The results of decoding the DNA sequence of 16 taxons of Anacardiaceae family in Vietnam that combined with information on Gen Bank built the relationship between taxons. At the same time, analyzing and explaining the scientific and objectivity of the traditional classification systems (morphological competition) about the Cashew family and classification systems of Engler (1892, 1896, 1903), Menchior (1964) and Heywood (1996) were selected to sort and classify taxons in the Cashew family in Vietnam. It is a logical choice, also the first time for this family in Vietnam. From that result, a phylogenetic tree in the family has been built. - The results of DNA sequencing have contributed to clarify the objectivity and accuracy of the traditional classification methods by morphological competition for taxons in the Cashew family. Specifically, the results also showed 4 branches equivalent to 4 tribus of the taxa of the Cashew family in Vietnam.

24

LIST OF THE THESIS RELATED PUBLICATIONS 1. Nguyen Xuan Quyen, Nguyen Van Ha, Tran Thi Phuong Anh, Nguyen The Cuong. Characteristic of some allergy included species of Anacardiaceae in Vietnam, Proceedings of the 6th National Scientific Conference on Ecology and Biological Resources: pp. 819-824, 2015, Publishing House of Natural Science and Technology. 2. Nguyen Xuan Quyen, Nguyen Van Ha, Tran Thi Phuong Anh, Nguyen The Cuong. Useful Anacardiaceae plants in Vietnam, Proceedings of the 6th National Scientific Conference on Ecology and Biological Resources: pp. 1201-1205, 2015, Publishing House of Natural Science and Technology 3. Nguyen Xuan Quyen, Tran Thi Phuong Anh, Nguyen The Cuong. Toxicodendron wallichii (Hook. f.) Kuntze, a new record of Ancardiaceae from Vietnam, Thai Journal of Botany, vol. 9, No. 1, January-June, ISSN 1906-7038: pp. 15-18, 2017. 4. Nguyen Xuan Quyen, Tran Thi Phuong Anh, Nguyen The Cuong. Morphological and taxonomic study on the genus Toxicodenron Mill. (Anacardiaceae R. Br.) in Vietnam, Proceedings of the 7th National Scientific Conference on Ecology and Biological Resources: pp. 340-344, 2017, Publishing House of Natural Science and Technology. 5. Nguyen Xuan Quyen, Tran Thi Phuong Anh, Nguyen The Cuong, Nguyen Thi Phuong Trang, Nguyen Thi Hong Mai, Hoang Le Tuan Anh. Molecular characteristic of Melanorrhoea usitata Wall. in Vietnam based on trnL and rbcL sequences, Proceedings of the 7th National Scientific Conference on Ecology and Biological Resources: pp. 1416-1420. 2017, Publishing House of Natural Science and Technology. 6. Nguyen Xuan Quyen, Tran Thi Phuong Anh, Nguyen The Cuong. Application of rbcL and trnL-trnF sequencing to study classification of cashew family (Anacardiaceae) in Vietnam, Science and technology Journal of Agriculture & Rural development, No. 14, ISSN 1859-4581: pp. 3-11; 2020.