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Systematics of the Thai Calophyllaceae and Hypericaceae with Comments on the Kielmeyeroidae (Clusiaceae)

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Systematics of the Thai Calophyllaceae and Hypericaceae with Comments on the Kielmeyeroidae (Clusiaceae) THAI FOREST BULL., BOT. 46(2): 162–216. 2018. DOI https://doi.org/10.20531/tfb.2018.46.2.08 Systematics of the Thai Calophyllaceae and Hypericaceae with comments on the Kielmeyeroidae (Clusiaceae) CAROLINE BYRNE1, JOHN ADRIAN NAICKER PARNELL1,2,* & KONGKANDA CHAYAMARIT3 ABSTRACT The Calophyllaceae and Hypericaceae are revised for Thailand and their relationships to the Clusiaceae and Guttiferae are briefly discussed. Thirty-two species are definitively recognised in six genera, namely: Calophyllum L., Kayea Wall., Mammea L. and Mesua L. in the Calophyllaceae and Cratoxylum Blume. and Hypericum L. in the Hypericaceae. A further four species of Calophyllum are tentatively noted as likely to occur in Thailand. Descriptions, full synonyms relevant to the Thai taxa, distribution maps, ecology, phenology, vernacular names, specimens examined and provisional keys are given. All species previously classified in the genus Mesua have been moved to the genus Kayea, except Mesua ferrea L. Two taxa were found to be endemic to Thailand: Mammea harmandii (Pierre) Kosterm. and Hypericum siamense N.Robson. The distribution for the families in Thailand was found to vary with the Thai Calophyllaceae being found mainly in Central and Peninsular Thailand whilst the Thai Hypericaceae were found mainly in the North and the North-East of Thailand. Overall the numbers of collections housed in herbaria are few and more collections are necessary in order to give a comprehensive account of their distributions in Thailand. KEYWORDS: Guttiferae, Flora of Thailand. Published online: 24 December 2018 INTRODUCTION from herbarium notes or directly from dried material. Ecological information was taken from specimens, The present work forms the basis of an account from field observations and from the literature. being prepared for the Flora of Thailand. It is largely Distribution maps were constructed using Arc-GIS based on the examination of 2,345 specimens from 9, Version 9.1. These maps show the distribution of worldwide herbaria including: A, AAU, ABD, B, all species in Thailand. The co-ordinates of localities BK, BKF, BM, C, CAL, CMUB, E, G, K, KKU, L, used have been taken from label data, and in the cases LINN, NY, P, PSU, QBG, SING, TCD, U and UPS. where only province names have been provided, a All descriptions are based on observations made co-ordinate point was mapped in the centre of that from herbarium material and living plants observed province to show the presence of the specimen. All while in the field. Dimensions used are based on types that have been seen are indicated by (!) after herbarium specimens and living plants measured in the herbarium abbreviation. Where a new lectotype the field. For dry specimens, flowers were softened has been chosen, this is indicated after the designated in water before measurements were taken. Vernacular specimen. All lectotypes have been chosen on the names and geographical information were taken same basis: that is that they are the best available of from specimens and the literature. Flower, and bark all potential lectotype specimens or, in rare cases, the exudate colours in Hypericum and Cratoxylum always sole available specimen. In the synonymy, homotypic refer to fresh material: all other colours are drawn 1 Herbarium, Botany Department, School of Natural Sciences, Trinity College Dublin, Dublin 2, Ireland. 2 Trinity Centre for the Environment, School of Natural Sciences, Trinity College Dublin, Dublin 2, Ireland. 3 Forest Herbarium, National Park, Wildlife and Plant Conservation Department, 61 Phahonyothin Road, Chatuchak, Bangkok 10900, Thailand. * Corresponding author: [email protected] © 2018 The Forest Herbarium SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE) (C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT) 163 synonyms are indicated by the identity sign (≡). Bar Clusioideae was sister to the Bonnetiaceae (Wurdack codes are given where possible; but, for example, & Davis, 2009). So as to ensure monophyletic family neither ABD or LINN use bar codes and in many circumscriptions Wurdack & Davis (2009) reinstated herbaria not all specimens are bar-coded. It is hoped the Calophyllaceae, a decision repeatedly and recently that this account will enable the keys, descriptions re-confirmed (APG IV, 2016). Sun et al. (2016) and distributional information on taxa to be refined undertook a mega-analysis of a super-matrix of 9,300 before they are published in the Flora of Thailand. species and confirmed that the Podostemaceae and Furthermore, it is clear that a number of taxa are Hypericaceae form a clade (100% Bootstrap support known from very few collections indeed, sometimes [BS]) with the Calophyllaceae being that clade’s only a single collection (e.g., Kayea elegans King): a sister (93% BS), this latter clade being sister to a situation repeatedly drawn attention to as a general more weakly supported clade (72% BS) comprising problem for many taxa in Thailand (Parnell et al., the Clusiaceae and Bonnetiaceae and with the whole 2003; van Welzen et al., 2011). lot forming a clade with 100% BS within the Malphigiales. Sun et al. (2016) also indicate that The Calophyllaceae and Hypericaceae and the Mesua in the Calophyllaceae is not monophyletic Clusiaceae and the Guttiferae necessitating the transfer of a number of species The families Calophyllaceae and Hypericaceae away from this genus. Some of these relationships are dealt with together in this paper as their status appear difficult to accept from a morphological as separate families has varied over time. Indeed, in perspective. In particular, the Podostemaceae are an the past, they have been dealt with as one family aquatic family of angiosperms that occur on rocks under the name Guttiferae or Clusiaceae, and that in waterfalls and rapids (Kato, 2004) and appear is how they were first proposed to be treated for the quite different to the other families in the Malpigiales Flora of Thailand when this work commenced. in terms of habit, indeed different to any other family of Angiosperm. Koi (2005) suggested that saltational The Clusiaceae, Calophyllaceae and evolution might have occurred to give rise to the Hypericaceae belong to the order Malpighiales Podostemaceae, as there is a lack of intermediate (Chase et al., 1993; Davis et al., 2004; Stevens, 2006) species between this family and the Clusiaceae s.l. which contains 700 genera and over 16,000 species and Hypericaceae. But there are morphological in 30 families (Tokuoka &Tobe, 2006; Wurdack & similarities linking them to each other, including Davis, 2009). The phylogenetic placement of this various types of secretory structure and secondary order is poorly understood; it may lie either in the products (Kubitzki et al., 1978; Stevens, 2006; fabid clade of the rosids (rosid I) or the malvid clade Wurdack & Davis, 2009). (rosid II) (APG IV, 2016). Phylogenetic relationships within the Malpighiales are also not fully resolved The Guttiferae (Clusiaceae, nom. alt.) sensu (Stevens, 2001 onwards; Davis et al., 2004; APG lato, comprises 27 genera and 1,050 species. Stevens II, 2003; Tokuoka & Tobe, 2006; APG III, 2009, (2006) considered there to be two subfamilies: Wurdack & Davis, 2009; Sun et al. 2016) though the Kielmeyeroideae and Clusioideae. interfamily relationships are now better understood The Kielmeyeroideae, now considered a separate (APG IV, 2016). Most recent phylogenetic studies family Calophyllaceae (Stevens, 2001 onwards; have been based on molecular analyses, using plastid APG III, 2009; APG IV, 2016; Mabberley, 2017) and nuclear gene regions (Savolainen et al., 2000a, contain two tribes, namely tribe Calophylleae with 2000b; Soltis et al., 2000; Stevens, 2001 onwards; 11 genera and tribe Endodesmieae with two genera APG II, 2003; Davis et al. 2004, 2005; Tokuoka & (Stevens, 2001 onwards). They are characterised by Tobe, 2006; Wurdack & Davis, 2009; APG III, 2009; spiral leaves with pellucid dots, moderate-sized APG IV, 2016; Sun et al. 2016). Wurdack & Davis cotyledons and perfect flowers and as Stevens (2006) (2009) showed that the Clusiaceae are polyphyletic indicates (as Clusiaceae) are found mainly in moist, with the two commonly recognised subfamilies of tropical, lowland or lower montane forests. The the Clusiaceae s.l. not forming a clade: subfamily majority of genera are found in primary forests, some Kielmeroideae was shown to be sister to the species grow in peat swamp forests (Calophyllum Hypericaceae and Podostemaceae, while subfamily 164 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2 pisiferum Planch. & Triana, C. rupicola Ridl., in the genus Mammea (Clusiaceae): Mammea C. sclerophyllum Vesque, C. teysmannii Miq.), and harmandii (Pierre) Kosterm. and M. siamensis (Miq.) some grow in black-water floodplains (species of T.Anderson. Three species were reported in the Caraipa Aublet, Haploclathra Benth.) (Stevens, genus Mesua (Clusiaceae), namely Mesua ferrea L., 2006). M. ferruginea (Pierre) Kosterm. and M. nervosa Planch. & Triana. The Clusioideae, now considered a separate family Clusiaceae (Stevens, 2001 onwards; APG III, 2009; APG IV, 2016; Mabberley, 2017), contain 14 TAXONOMIC ACCOUNT genera and 500 species (Mabberley, 2017) in three The following is a brief description of the four tribes (Stevens, 2001 onwards), namely Clusieae genera and 21(–25) species of Calophyllaceae and with five genera, Garcinieae with two generan and the two genera and 12 species of Hypericaceae that Symphonieae with seven genera, and further details we
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