Garcinia section Brindonia (Thouars) Choisy, Mem . Soc. Hist. Nat. Paris 1: 230 (1823). Clade 5

Gaudeul, Myriam, Sweeney, Patrick & Munzinger, Jerome, 2024, An updated infrageneric classification of the pantropical species-rich genus Garcinia L. (Clusiaceae) and some insights into the systematics of New Caledonian species, based on molecular and morphological evidence, PhytoKeys 239, pp. 73-105 : 73

6. Garcinia section Brindonia (Thouars) Choisy, Mem. Soc. Hist. Nat. Paris 1: 230 (1823). Clade 5

Figs 2 , 3

Basionym.

Brindonia Thouars, Dict. Sci. Nat. [F. Cuvier] 5: 339 (1806).

Type.

Brindonia oxycarpa Thouars, Hist. Veg. Isles Austr. Afr. Ed. 2 t. 27 (1805) [≡ Garcinia oxycarpa (Thouars) P.W.Sweeney, comb. nov.; Garcinia indica (Thours) Choisy Mém. syn. nov.]. The copy of Histoire des végétaux recueillis dans les isles australes d’Afrique ed. 2 at Kew bears the date 1805 ( Baker 1894) and contains six plates (25-30) that do not have accompanying text in the main body of the publication (see also Hiern 1900). Plate 27 is labeled " Brindonia oxycarpa " and it depicts two flowering branches, fruit, and dissected flowers (pers. obs). This suffices as an illustration with analysis and thus Brindonia oxycarpa is validly published as per Articles 38.1, 38.7, and 38.8 of the ICN ( Turland et al. 2018). Some (e.g. Hiern 1900) have considered B. oxycarpa a synonym of Garcinia indica (Thours) Choisy Mém. Soc. Hist. Nat. Paris 1: 230 (1823) [≡ Brindonia indica Thouars, Dict. Sci. Nat. [F. Cuvier] 5: 340 (1806)]; however, if these two taxa are considered synonymous, the epithet oxcycarpa would have priority.

Distinguishing sectional characters.

Flowers with four sepals and petals. Staminate flowers without a pistillode (usually), stamens united into a single central bundle (or ring when pistillode present), anthers four-thecous (but in some species two-thecous). Ovaries multilocular, stigmas divided into distinct rays and usually papillate. Fruits in many species with furrows or grooves along the septal radii. Inflorescence s terminal or axillary with one to many flowers. Afrotropics (Madagascar), Indomalaya, tropical Australasia, and Oceania.

Three species treated as Garcinia Garcinia by Jones (1980) (i.e., G. costata Hemsl. ex King, G. maingayi Hook.f., and G. trianii Pierre) form a clade sister to clade 5, the latter largely corresponding to Garcinia Brindona sensu Jones (1980). While they share some features (e.g. tendency to have furrowed/grooved (very shallow in G. maingayi and G. trianii ), multilocular fruits) with Brindonia sensu Jones (1980), they have other features (i.e., pistillodes, stamens arranged into a ring, and two-thecous anthers) that are not typical of the section. While it would be tempting to recognize a separate section for these species, pistillodes and stamens arranged into a ring are also shared by G. atroviridis Griff. ex T.Anderson and G. pedunculata Roxb. ex Buch.-Ham., two species that were included in Garcinia Brindonia by Jones (1980). It is noteworthy that G. atroviridis is the first branching lineage within clade 5, which together with the G. costata / G. maingayi / G. trianii clade form a basal grade within Garcinia Brindonia (as circumscribed here).

Garcinia usually has an indehiscent drupe or berry ( Stevens 2007). The genus Clusianthemum Vieill. was established by Vieillard for a new Caledonian species having a capsular fruit ( C. pedicellatum Vieill.). Later, another capsular genus, Septogarcinia was established by Kostermans (1962) for S. sumbawaensis Kosterm., from Sumbawa (Indonesia), obviously not knowing about Vieillard’s Clusianthemum . Notably, several species of NC Garcinia , viz. G. virgata Vieill. ex Guillaumin, G. neglecta Vieill. and G. comptonii Baker f. have capsular fruits (Fig. 1 ). Jones (1980) did not mention Clusianthemum in her treatment but included Septogarcinia in Garcinia sect. Brindonia . Jones (1980) does not cite any NC capsular species. The newly described G. urceolata is also suspected of having dehiscent fruits ( Munzinger et al. 2021). All these species are morphologically similar and could result from in situ (within NC) diversification, and all sampled species with dehiscent fruit are found in a strongly supported subclade within clade 5. We do not have sequence material of G. septogarcinia I.M. Turner & L.V.S. Jenn. to determine whether that character is an autapomorphy of a dehiscent fruit clade, or if it evolved at least twice, in New Caledonia and Sumbawa. Staminate floral morphology supports placement of G. septogarcinia I.M. Turner & L.V.S. Jenn. into Garcinia Brindona ( Medellín-Zabala and Marinho 2015).

Species.

Garcinia adinantha A.C.Sm. & S.P.Darwin; G. amabilis Kaneh. & Hatus.; G. amboinensis Spreng.; G. angustifolia A.C. Sm.; G. assamica J.Sarma, Shameer & N.Mohanan; G. assugu Lauterb.; G. asterandra Jum. & H.Perrier; G. atroviridis Griff. ex T.Anderson; G. balimensis A.C. Sm.; G. bancana Miq.; G. beccarii Pierre; G. bicolorata Elmer; G. binucao (Blanco) Choisy; G. borneensis Pierre; G. chapelieri (Planch. & Triana) H.Perrier; G. cochinchinensis (Lour.) Choisy; G. comptonii Baker f.; G. conicarpa Wight; G. corallina Vieill.; G. costata Hemsl. ex King; G. cowa Roxb. ex DC.; G. crassiflora Jum. & H.Perrier; G. dallmannensis Kaneh. & Hatus.; G. delpyana Pierre; G. dhanikhariensis S.K.Srivast.; G. dioica Blume; G. emarginata Lauterb.; G. erythrosepala Y.H.Li; G. esculenta Y.H.Li; G. fruticosa Lauterb.; G. fusca Pierre; G. griffithii T.Anderson; G. gummi-gutta (L.) N.Robson; G. horsfieldiana Pierre; G. hygrophila Lauterb.; G. indica (Thouars) Choisy; G. klinkii Lauterb.; G. korthalsii Pierre; G. lanceifolia Roxb.; G. lauterbachiana A.C.Sm.; G. ledermannii Lauterb.; G. leggeae W.E.Cooper; G. loheri Merr.; G. macgregorii Merr.; G. macrantha A.C.Sm.; G. maingayi Hook. f.; G. maluensis Lauterb.; G. mestonii F.M.Bailey; G. microstigma Kurz; G. minahassensis Pierre; G. miquelii Pierre; G. myristicifolia Pierre; G. nigrolineata Planch. ex T.Anderson; G. oblongifolia Champ. ex Benth.; G. oligophlebia Merr.; G. oliveri Pierre; G. oreophila Lauterb.; G. oxycarpa (Thours) P.W.Sweeney; G. pachyantha A.C.Sm.; G. pachypetala Lauterb.; G. pallida Lauterb.; G. parvifolia (Miq.) Miq.; G. pedunculata Roxb. ex Buch.-Ham.; G. ponapensis Lauterb.; G. quaesita Pierre; G. ramosii Merr.; G. riparia A.C.Sm.; G. rubra Merr.; G. rubriflora Boerl.; G. sabangensis Lauterb.; G. samarensis Merr.; G. schomburgkiana Pierre; G. segmentata Kosterm.; G. septogarcinia I.M.Turner & L.V.S.Jenn.; G. siripatanadilokii Ngerns., Meeprom, Boonth., Chamch. & Sinbumr.; G. solomonensis A.C.Sm.; G. sopsopia (Buch.-Ham.) Mabb.; G. stigmacantha Pierre; G. succifolia Kurz; G. sulphurea Elmer; G. tetrandra Pierre; G. teysmanniana Scheff.; G. trianii Pierre; G. urceolata Munzinger, Bruy & M.Pignal; G. valetoniana Lauterb.; G. vidua Ridl.; G. virgata Vieill. ex Guillaumin; G. viridiflora Ridl.; G. wallichii Choisy; G. xishuanbannaensis Y.H.Li; G. zeylanica Roxb.

Figures

Figure 1. Some Garcinia New Caledonian species (except E from Fiji) and morphological features A G. balansae (Munzinger 4916), fruiting branch B G. balansae (Munzinger 4916), bark C G. sp. " JT 814 " (Munzinger 7282), habit D G. sp. " JT 814 " (Munzinger 7282), bark E G. vitiensis (Munzinger 7377), fruiting branch F G. neglecta (Munzinger 2690), fruit G G. comptonii (sin voucher), fruit.

Figure 2. Molecular phylogeny of Garcinia L. based on ITS sequences and Bayesian inference. Posterior probabilities (PP) and bootstrap support values (BS), obtained respectively by the Bayesian inference and Maximum Likelihood (ML) analysis, are indicated at each node of the cladogram. Nodes were collapsed when PP <0.50. The lineages / sections discussed in the text are highlighted, and species names appear in colors depending on their native distribution areas: light blue, Central and South America; light green, Tropical Africa; dark green, Madagascar and Western Indian Ocean islands; grey, Southeast Asia; purple, Australia; orange, New Guinea; red, New Caledonia; dark blue, Southwest Pacific islands. Distribution information was taken from the Plants of the World Online website (POWO 2023; also see the table of vouchers). A few species occur in several regions, and the color of the main (largest) geographic region was used. Accessions in bold were newly sequenced in this study.

Figure 3. Molecular phylogeny of Garcinia L. based on a combined chloroplast DNA dataset and Bayesian inference. Posterior probabilities (PP) and bootstrap support values (BS), obtained respectively by the Bayesian inference and Maximum Likelihood (ML) analysis, are indicated at each node of the cladogram. Nodes were collapsed when PP <0.50. The lineages / sections discussed in the text are highlighted, and species names appear in colors depending on their native distribution areas: light green, Tropical Africa; dark green, Madagascar and Western Indian Ocean islands; grey, Southeast Asia; purple, Australia; orange, New Guinea; red, New Caledonia; dark blue, Southwest Pacific islands. Distribution information was taken from the Plants of the World Online website (POWO 2023; also see the table of vouchers). A few species occur in several regions, and the color of the main (largest) region was used. All accessions were newly sequenced in this study.