Research Article |
Corresponding author: Richard M. K. Saunders ( saunders@hku.hk ) Academic editor: Thomas L.P. Couvreur
© 2021 Charan Leeratiwong, Piya Chalermglin, Richard M. K. Saunders.
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:
Leeratiwong C, Chalermglin P, Saunders RMK (2021) Goniothalamus roseipetalus and G. sukhirinensis (Annonaceae): Two new species from Peninsular Thailand. PhytoKeys 184: 1-17. https://doi.org/10.3897/phytokeys.184.73210
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Two new Goniothalamus species (Annonaceae), G. roseipetalus sp. nov. and G. sukhirinensis sp. nov., are described from the southern limits of Peninsular Thailand (Narathiwat and Yala Provinces). Both new species resemble G. macrophyllus, G. scortechinii and G. uvarioides. The addition of these two new species brings the total number of Goniothalamus species in Thailand to 27. Separate identification keys are provided for flowering and fruiting specimens of the Thai species.
Annonaceae, Goniothalamus roseipetalus, Goniothalamus sukhirinensis, new species, Thailand
The genus Goniothalamus (Blume) Hook.f. & Thomson (Annonaceae subfam. Annonoideae tribe Annoneae:
Goniothalamus fruits are apocarpous, with distinct fleshy ‘monocarps’ that develop from individual carpels after fertilisation. Two contrasting seed dispersal systems have been inferred, correlated with differences in fruit and seed morphology (
Goniothalamus is comparatively species-rich, with over 130 species. Although the genus has never been comprehensively revised, there are several recent regional taxonomic studies, including Thailand (
Goniothalamus roseipetalus resembles G. scortechinii and G. uvarioides but is distinguished by its leaves with generally fewer secondary veins (15–22 pairs), wider sepals (24–35 mm), and wider inner petals (8–11 mm). It is also distinguished from G. scortechinii by its wider outer petals (14–25 mm), and is distinguished from G. uvarioides by its smaller, single-seeded monocarps (8–15 by 7–9 mm), borne on shorter stipes (3–5 mm).
Goniothalamus roseipetalus sp. nov. A flowering branch B flower C calyx of fused sepals (abaxial) D calyx of fused sepals (adaxial) E outer petal (abaxial) F outer petals (adaxial) G inner petal (abaxial). H inner petal (adaxial) I stamen (abaxial) J stamen (adaxial) K carpel (abaxial) L carpel (adaxial) M fruit, composed of separate monocarps. N–P seeds (different orientations). Drawn by A. Somphrom A–L from C. Leeratiwong 21–1708 (PSU) M–P from C. Leeratiwong 21–1707 (PSU).
Shrubs to small trees, to 4 m. Young branches glabrous. Leaf laminas 15–40 by 3–13 cm, length/width ratio 2.8–5, elliptic to oblanceolate, apex generally acuminate (rarely acute to obtuse), acumen 3–10 mm long, base broadly cuneate, chartaceous, glabrous ab- and adaxially (sometimes sparsely pubescent over midrib); midrib strongly prominent abaxially, sunken adaxially; secondary veins 15–22 pairs, plane adaxially; tertiary veins percurrent, slightly distinct, lacking a ‘granular’ appearance abaxially; petioles 12–22 mm by 1.5–2.5 mm, glabrous to sparsely pubescent. Flowers solitary, often on main trunk (cauliflorous), rarely on older branches (ramiflorous), pendent; flowering pedicels 10–17 mm long, sparsely hairy; pedicel bracts ovate to broadly lanceolate, 2–4 by 2–3 mm. Sepals (violet-)pink, broadly ovate, 20–30 by 24–35 mm, basally connate (10–17 mm from base), apex rounded, glabrous ab- and adaxially, with sparsely hairy margins, venation distinct, 5–7-veined. Outer petals greenish-pink when young, (violet-)pink (green at claw) when mature, 25–45 by 14–25 mm with 4–10 mm-long claw, length/width ratio 1.7–2.2, fleshy, (lanceolate-)ovate, apex obtuse to mucronate, reflexed, sparsely hairy abaxially (more densely along margins basally), sparsely hairy (more densely apically) adaxially with velutinous basal region facing apertures between inner petals, midrib and venation indistinct ab- and adaxially. Inner petals 12–20 by 8–11 mm with 2–5 mm-long claw, length/width ratio 1.5–1.8, oblanceolate, densely hairy ab- and adaxially, greenish-pink when young, pale pink when mature, apex acute, lacking a glabrous lasteral flange on the inner petal claws. Stamens numerous, narrowly oblong, 3–4 mm long; connectives apiculate, papillate. Carpels 20–35 per flower, ovary oblong, 2–2.5 mm long, with white hairs; stigma and pseudostyle 2–3 mm long, stigma subulate, glabrous. Fruits with persistent calyx, immature fruits greenish-pink, mature fruits (pinkish-)red; fruiting pedicels 10–20 by 2–2.5 mm, sparsely hairy to glabrous. Monocarps 5–20 per fruit, 1–2-seeded, 8–17 by 7–10 mm, length/width ratio 1.1–1.7, ellipsoid to ovoid, apex apiculate, apicule 0.5–1.5 mm long, smooth, sparsely hairy, glossy, pericarp 1–2 mm thick, stipes 3–6 by 1.5–2 mm, moderately hairy. Seeds with mucilage, 9–11 by 8–9 mm, length/width ratio 1.1–1.6, ovoid, testa sparsely pubescent, rugose.
Flowering in March and April; fruiting in August (based on limited data).
Endemic to Peninsular Thailand, where it occurs in Narathiwat and Yala Provinces (Fig.
In reference to the red pigmentation of the petals.
Panan klip muang (ปาหนันกลีบม่วง) (general).
Thailand: Yala Province, Bannang Sata, 350 m alt., 1 August 2020, C. Leeratiwong 20–1684 (PSU); Narathiwat Province: Cha Nae District, Du Son Yo subdistrict, 100 m alt., 6 March 2021, C. Leeratiwong 21–1705 (PSU).
Although G. roseipetalus is yet to be included in a molecular phylogenetic analysis, it shares several morphological similarities with species in a clade (nested within clade ‘A1a’ sensu
Goniothalamus roseipetalus is morphologically most similar to G. scortechinii and G. uvarioides. It differs from these species, however, as it generally has fewer secondary veins in its leaves (15–22 pairs, vs [18–]21–26[–32] in G. scortechinii and 24–35 in G. uvarioides), larger sepals (20–30 by 24–35 mm, vs 8–24 by 8–23 mm in G. scortechinii and 12–16 by 5–13 mm in G. uvarioides), and wider inner petals (8–11 mm, vs 5–8 mm in G. scortechinii and 7–8.5 mm in G. uvarioides). It also has wider outer petals (14–25 mm) than G. scortechinii (8–14 mm), and can be distinguished from G. uvarioides by reference to its smaller monocarps (8–15 by 7–9 mm, vs 31–44 by 15–18 mm) with a single seed (vs four or five seeds per monocarp) and shorter stipes (3–5 mm, vs 12.5–17.5 mm). Goniothalamus roseipetalus also resembles the widespread species G. macrophyllus, although the latter species has creamy-white petals.
Goniothalamus roseipetalus also resembles G. calycinus J. Sinclair, a species that is endemic to Terengganu in Peninsular Malaysia (
Goniothalamus sukhirinensis resembles G. macrophyllus and G. scortechinii, but is distinguished by its densely hairy shoots, numerous secondary veins (32–40 pairs per leaf), generally longer pedicels (flowering: 12–18 mm; fruiting: 20–25 mm), larger outer petals (34–37 by 18–22 mm), larger monocarps (20–27 by 9–13 mm) that are densely hairy, and longer seeds (13–17 mm).
Goniothalamus sukhirinensis sp. nov. A vegetative branch B petiole, showing base of leaf lamina C flowers D sepal (abaxial) E sepal (adaxial) F outer petal (abaxial) G outer petals (adaxial) H inner petal (abaxial) I inner petal (adaxial) J stamen (abaxial) K stamen (adaxial) L carpel (abaxial) M carpel (adaxial) N fruit, composed of separate monocarps O seed with hairy surface. Drawn by A. Somphrom from C. Leeratiwong 21–1708 (PSU).
Shrubs to small trees, to 4 m. Young branches densely appressed-pubescent. Leaf laminas 28–50 by 7–16 cm, length/width ratio 3.1–4, (lanceolate-)oblong, apex generally acuminate to caudate (rarely acute to obtuse), acumen 7–20 mm long, base broadly cuneate, subcoriaceous, glabrous abaxially (sparsely hairy over midrib), sparsely pubescent adaxially (densely hairy over veins); midrib strongly prominent abaxially, sunken adaxially; secondary veins 32–40 pairs, plane adaxially; tertiary veins percurrent, distinct, lacking a ‘granular’ appearance abaxially; petioles 20–30 mm by 4–6 mm, densely pubescent. Flowers solitary or paired, often on main trunk (cauliflorous), rarely on older branches (ramiflorous), pendent; flowering pedicels 12–18 mm long, densely hairy; pedicel bracts ovate-triangular, 2.5–3 by 1–1.5 mm. Sepals greenish-pink, broadly ovate, 7–9.5 by 7.5–10 mm, basally connate (2.5–3 mm from base), apex acute, moderately hairy abaxially, sparsely hairy adaxially, venation indistinct. Outer petals greenish-yellow when young, whitish-yellow (green at claw) when mature, 34–37 by 18–22 mm with 3–5 mm-long claw, length/width ratio 1.6–1.9, fleshy, (lanceolate-)ovate, apex acuminate, densely hairy abaxially, moderately hairy adaxially with velutinous basal region facing apertures between inner petals, midrib raised adaxially, venation indistinct ab- and adaxially. Inner petals 13–15 by 7–8 mm with 2–3 mm long claw, length/width ratio 1.8–1.9, ovate-lanceolate, densely hairy abaxially, sparsely hairy distally adaxially, yellowish-green when young, pinkish-orange to reddish-brown when mature, apex acuminate, lacking a glabrous lasteral flange on the inner petal claws. Stamens numerous, oblong, 2.5–3.7 mm long; connectives apiculate, papillate. Carpels 11–20 per flower, ovary oblong, 2–3 mm long, with white hairs; stigma and pseudostyle 2–2.5 mm long, stigma funnel-shaped, hairy. Fruits sometimes with persistent calyx, immature fruits brownish-green, mature fruits not seen; fruiting pedicels 20–25 by 2–3.5 mm, sparsely hairy. Monocarps 5–14 per fruit, single-seeded, 20–27 by 9–13 mm, length/width ratio 2–2.7, (obovoid-)ellipsoid, apex apiculate, apicule 5–8 mm long, smooth, densely hairy, glossy, pericarp 1–2 mm thick, stipes 7–15 by 2–3 mm, densely hairy. Seeds 13–17 by 8–10 mm, length/width ratio 1.6–1.7, ellipsoid, testa densely villose, slightly rugose.
Flowering and fruiting in February and March (based on limited data).
Endemic to Narathiwat Province, Peninsular Thailand (Fig.
From the name Sukhirin, Narathiwat Province.
Ratchakhru khao (ราชครูขาว) (Narathiwat).
Thailand: Narathiwat Province: Sukhirin District, Ban Yade village, Ma Mong subdistrict, 200 m alt., 28 February 2021, C. Leeratiwong 21–1707 (PSU).
As with the previous species, G. sukhirinensis is yet to be included in a molecular phylogenetic analysis but has strong morphological affiliations with a clade that comprises G. loerzingii R.M.K.Saunders, G. macrophyllus, G. scortechinii, G. uvarioides and G. wrayi King (nested within clade ‘A1a’ sensu
Goniothalamus sukhirinensis resembles G. macrophyllus and G. scortechinii, but differs in several key characters: densely hairy shoots (vs glabrous to medium-hairy); numerous secondary veins (32–40 pairs per leaf, vs 12–23 in G. macrophyllus and [18–]21–26[–32] in G. scortechinii); generally longer flowering pedicels (12–18 mm, vs 5–11.5 mm in G. macrophyllus and 8–13 mm in G. scortechinii); larger outer petals (34–37 by 18–22 mm, vs 10–28 by 4.5–11.5 mm in G. macrophyllus and 20–33 by 8–14 mm in G. scortechinii); longer fruiting pedicels (20–25 mm, vs 7–19 in G. macrophyllus and 8–20 mm in G. scortechinii); larger monocarps (20–27 by 9–13 mm, vs 8–15 by 7.5–10 mm in G. macrophyllus and 9–18 by 6–10 mm in G. scortechinii) that are densely hairy (vs subglabrous to medium-hairy); and longer seeds (13–17 mm, vs 8.5–12 mm in G. macrophyllus and 8–11 mm in G. scortechinii). Goniothalamus sukhirinensis also differs from G. macrophyllus as its leaves lack the fine ‘granular’ appearance of the latter species (due to the immersion of tertiary and higher-order veins: Saunders, 2002), and has longer monocarp stipes (7–15 mm, vs up to 1.8 mm in G. macrophyllus).
1a | Stamen connective apex apiculate | 2 |
2a | Young branches densely hairy to velutinous | 3 |
3a | Leaf laminas 28–50 cm long, with 32–40 pairs of secondary veins; sepals 7–9.5 by 7.5–10 mm; outer petals 34–37 by 18–22 mm; inner petals 13–15 by 7–8 mm | G. sukhirinensis sp. nov. |
3b | Leaf laminas 50–76 cm long, with 24–32 pairs of secondary veins; sepals 30–40 by 28–30 mm; outer petals 60–80 by 30–40 mm; inner petals ca. 35 by ca. 17 mm | G. cheliensis H.H.Hu |
2b | Young branches glabrous to hairy | 4 |
4a | Flowers in large fascicles, exclusively from woody tubercles at base of trunk | G. ridleyi King |
4b | Flowers solitary or in pairs, not exclusively from base of trunk | 5 |
5a | Adaxial surface of outer petals with glabrous or sparsely hairy region facing apertures between inner petals | 6 |
6a | Leaves with 24–35 pairs of secondary veins | G. uvarioides King |
6b | Leaves with 11–22 pairs of secondary veins | G. tapis Miq. |
5b | Adaxial surface of outer petals with velutinous region facing apertures between inner petals | 7 |
7a | Flowering pedicels 20–37 mm long; carpels 50–100 per flower | G. tortilipetalus M.R.Hend. |
7b | Flowering pedicels 5–19 mm long; carpels 8–50 per flower | 8 |
8a | Leaves with 9–12 pairs of secondary veins; stamen connective apex distinctly tapered; carpels 8–10 per flower | G. tavoyensis Chatterjee |
8b | Leaves with 12–26(–32) pairs of secondary veins; stamen connective apex not distinctly tapered; carpels 11–50 per flower | 9 |
9a | Tertiary venation reticulate | 10 |
10a | Sepals 11–18.5 mm long, 8–15.5 mm wide; outer petals 21–46 mm long, 4.5–18 mm wide, yellow; inner petal length/width ratio 3–6.5; stamens 95–120 per flower | G. calvicarpus Craib |
10b | Sepals 14–29 mm long, 12–26 mm wide; outer petals 36–104 mm long, 14–24 mm wide, green; inner petals length/width ratio 1.8–3.6; stamens 100–200 per flower | G. griffithii Hook.f. & Thomson |
9b | Tertiary venation percurrent | 11 |
11a | Sepals 24–35 mm wide; outer petals 14–25 mm wide | G. roseipetalus sp. nov. |
11b | Sepals 4–23 mm wide; outer petals 4.5–14 mm wide | 12 |
12a | Leaf laminas (sub-)coriaceous, with fine “granular” texture abaxially (due to immersion of tertiary and lower order veins); leaves with 12–23 pairs of secondary veins | G. macrophyllus (Blume) Hook.f. & Thomson |
12b | Leaf laminas papyraceous, without fine ‘granular’ texture abaxially; leaves with (18–)21–26(–32) pairs of secondary veins | G. scortechinii King |
1b | Stamen connective apex truncate | 13 |
13a | Inner petal claws with distinct glabrous lateral flange | 14 |
14a | Flowering pedicels 7–23 mm long; stigma subulate | 15 |
15a | Outer petals 23–43 mm long, 12–23 mm wide; carpels 40–100 per flower | G. sawtehii C.E.C.Fisch. |
15b | Outer petals 10.5–32 mm long, 5.5–17.5 mm wide; carpels 10–54 per flower | G. undulatus Ridl. |
14b | Flowering pedicels 2–11.5 mm long; stigma fusiform or funnel-shaped | 16 |
16a | Young branches densely hairy to velutinous | G. tamirensis Pierre ex Finet & Gagnep. |
16b | Young branches glabrous to hairy | 17 |
17a | Flowering pedicels 5–11.5 mm long; sepal venation generally indistinct; outer petals 12.5–73 mm long; stigma fusiform | G. laoticus (Finet & Gagnep.) Bân |
17b | Flowering pedicels 2–6 mm long; sepal venation distinct; outer petals 8.5–39 mm long; stigma funnel-shaped | 18 |
18a | Leaf laminas 8–14 cm long, 1.5–4 cm wide; petioles 3.5–7 mm long; sepals 3–9 mm long, 3.5–6 mm wide; outer petals 8.5–15 mm long, 3.5–8 mm wide, very densely hairy ab- and adaxially; inner petals 6.5–10 mm long, 3–4.5 mm wide; ovary glabrous | G. elegans Ast |
18b | Leaf laminas 12.5–24.5 cm long, 4–8.5 cm wide; petioles 5–15 mm long; sepals 7.5–12.5 mm long, 5.5–11 mm wide; outer petals 23–39 mm long, 7–15 mm wide, glabrous to hairy ab- and adaxially; inner petals 10–16 mm long, 5.5–9 mm wide; ovary sparsely hairy | G. latestigma C.E.C.Fisch. |
13b | Inner petal claws without glabrous lateral flange | 19 |
19a | Tertiary leaf venation generally reticulate; outer petals with velutinous region at base of adaxial surface (facing aperture between inner petals); inner petals velutinous adaxially | 20 |
20a | Sepals 2.5–3.5 mm long, basally connate, venation indistinct; outer petal venation distinct; stigma fusiform | G. repevensis Pierre ex Finet & Gagnep. |
20b | Sepals 4.5–19 mm long, free, venation distinct; outer petal venation indistinct; stigma subulate or funnel-shaped | 21 |
21a | Young branches glabrous; outer petal length/width ratio 3.4–5.2 | G. expansus Craib |
21b | Young branches very sparsely to densely hairy; outer petal length/width ratio 1.6–3.8 | 22 |
22a | Leaves with fine “granular” texture abaxially (due to immersion of tertiary and lower order veins); flowers slightly supra-axillary | G. rotundisepalus M.R.Hend. |
22b | Leaves without fine “granular” texture abaxially; flowers axillary | G. tenuifolius King |
19b | Tertiary leaf venation percurrent; outer petals with glabrous or sparsely hairy region at base of adaxial surface (facing aperture between inner petals); inner petals glabrous to densely hairy adaxially | 23 |
23a | Flowering pedicels 20–48 mm long; sepals 7–15 mm long; outer petals 68–113 mm long, 28–63 mm wide | G. giganteus (Wall. ex) Hook.f. & Thomson |
23b | Flowering pedicels 8–16(–21) mm long; sepals 2–8 mm long; outer petals 16–50(–62) mm long, 7–22(–32) mm wide | 24 |
24a | Inner petals glabrous adaxially; ovaries densely hairy | G. malayanus Hook.f. & Thomson |
24b | Inner petals (densely) hairy adaxially (sometimes glabrous towards base); ovaries glabrous to sparsely hairy | 25 |
25a | Leaf laminas 23–32 cm long, with 14–21 pairs of secondary veins; stamens 50–160 per flower; carpels 4–11 per flower | G. aurantiacus R.M.K.Saunders & Chalermglin |
25b | Leaf laminas 17–25.5 cm long, with 13–16 pairs of secondary veins; stamens ca. 180–200 per flower; carpels ca. 18–20 per flower | 26 |
26a | Flowering pedicels densely hairy; outer petals densely hairy abaxially, very densely hairy adaxially, venation indistinct; inner petals very densely hairy abaxially | G. maewongensis R.M.K.Saunders & Chalermglin |
26b | Flowering pedicels very sparsely hairy; outer petals subglabrous abaxially, glabrous adaxially, venation distinct; inner petals sparsely hairy abaxially | G. rongklanus R.M.K.Saunders & Chalermglin |
1a | Adaxial surface of leaves with very prominent secondary veins | 2 |
2a | Leaf laminas 50–76 cm long, 13–22 cm wide, with 24–32 pairs of secondary veins; leaf midrib densely hairy to velutinous; petioles 17–30 mm long, velutinous; monocarps densely hairy | G. cheliensis H.H.Hu |
2b | Leaf laminas 12.5–39.5 cm long, 3.5–9.5(–11.5) cm wide, with 10–22 pairs of secondary veins; leaf midrib glabrous to sparsely hairy; petioles 4–16 mm long, glabrous to hairy; monocarps glabrous to hairy | 3 |
3a | Monocarps distinctly warty | G. giganteus (Wall. ex) Hook.f. & Thomson |
3b | Monocarps smooth or finely rugulose | 4 |
4a | Fruits restricted to trunk; fruiting pedicels 19–36 mm long | G. tortilipetalus M.R.Hend. |
4b | Fruits not restricted to trunk; fruiting pedicels 10–19 mm long | 5 |
5a | Tertiary leaf venation percurrent; fruits without persistent calyx; monocarps 16–40 mm long, 8–13(–17) mm wide; seeds 13–20 mm long, with (sparsely) hairy testa | G. malayanus Hook.f. & Thomson |
5b | Tertiary leaf venation reticulate; fruits with persistent calyx; monocarps 10–14 mm long, 7–8 mm wide; seeds 10–12 mm long, with glabrous testa | 6 |
6a | Monocarps red; seeds slightly rugose | G. calvicarpus Craib |
6b | Monocarps yellow-brown; seeds smooth | G. griffithii Hook.f. & Thomson |
1b | Adaxial surface of leaves with impressed or only slightly prominent secondary veins | 7 |
7a | Tertiary leaf venation percurrent | 8 |
8a | Monocarps apiculate | 9 |
9a | Young branches densely hairy; leaves with 32–40 pairs of secondary veins; fruiting pedicels 20–25 mm long; monocarps 20–27 mm long, densely hairy; monocarp stipes 7–15 mm long | G. sukhirinensis sp. nov. |
9b | Young branches glabrous to hairy; leaves with 12–23 pairs of secondary veins; fruiting pedicels 7–20 mm long; monocarps 8–15 mm long (very) sparsely hairy; monocarp stipes up to 5 mm long | 10 |
10a | Fruits without persistent calyx | G. macrophyllus (Blume) Hook.f. & Thomson |
10b | Fruits with persistent calyx | G. roseipetalus sp. nov. |
8b | Monocarps not apiculate | 11 |
11a | Monocarps 7–18 mm long, 6–10 mm wide | 12 |
12a | Leaf laminas 22–40(–50) cm long, 5.5–12(–19.5) cm wide, with (18–)21–26(–32) pairs of secondary veins | G. scortechinii King |
12b | Leaf laminas 8–20 cm long, 2.5–6 cm wide, with 8–14 pairs of secondary veins | G. tenuifolius King |
11b | Monocarps 16–56 mm long, 13–29 mm wide | 13 |
13a | Leaf laminas 32–48 cm long, 7–15.5 cm wide, with 24–35 pairs of secondary veins; fruits without persistent calyx | G. uvarioides King |
13b | Leaf laminas 14–32 cm long, 3.5–9.5 cm wide, with 11–21 pairs of secondary veins; fruits with persistent calyx | 14 |
14a | Fruits restricted to base of trunk; fruiting pedicels 30–130 mm long | G. ridleyi King |
14b | Fruits not restricted to base of trunk; fruiting pedicels 10–22 mm long | 15 |
15a | Monocarps subsessile or with stipe ≤ 1 mm long; seeds 18–19 mm wide, rugose | G. maewongensis R.M.K.Saunders & Chalermglin |
15b | Monocarp stipes 2–8 mm long; seeds 13–17 mm wide; smooth to slightly rugulose | 16 |
16a | Leaf laminas 23–32 cm long, with 14–21 pairs of secondary veins; fruiting pedicels ca. 14 mm long; seeds 19–27 mm long, length/width ratio 1.5–1.7 | G. aurantiacus R.M.K.Saunders & Chalermglin |
16b | Leaf laminas 17–25 cm long, with 13–16 pairs of secondary veins; fruiting pedicels 16–22 mm long; seeds 15–22 mm long, length/width ratio 1.1–1.5 | G. rongklanus R.M.K.Saunders & Chalermglin |
7b | Tertiary leaf venation reticulate | 17 |
17a | Monocarps 22–56 mm long, 14–21 mm wide, with prominent longitudinal ridge; up to 7 seeds per monocarp | G. laoticus (Finet & Gagnep.) Bân |
17b | Monocarps 8.5–26 mm long, 6–11 mm wide, without longitudinal ridge; 1 or 2 seeds per monocarp | 18 |
18a | Fruiting pedicels 8–20 mm long | 19 |
19a | Young branches (very densely) hairy | 20 |
20a | Leaf laminas glossy adaxially; monocarp stipes 6.5–16.5 mm long; seeds 9–11.5 mm long | G. undulatus Ridl. |
20b | Leaf laminas matt adaxially; monocarp stipes 3–6 mm long; seeds 11–14 mm long | G. sawtehii C.E.C.Fisch. |
19b | Young branches glabrous to hairy | 21 |
21a | Leaf lamina tertiary venation (clearly) distinct | G. expansus Craib |
21b | Leaf lamina tertiary venation indistinct to ± distinct | 22 |
22a | Leaf laminas with 9–12 pairs of secondary veins; monocarps greenish-yellow, ca. 19 mm long; seeds 17–18 mm long | G. tavoyensis Chatterjee |
22b | Leaf laminas with 11–16 pairs of secondary veins; monocarps (dark) red, 10–14 mm long; seeds 9–12 mm long | G. tapis Miq. |
18b | Fruiting pedicels 4–8 mm long | 23 |
23a | Monocarps greenish-yellow | 24 |
24a | Young branches hairy; leaf laminas 12.5–24.5 cm long, 4–8.5 cm wide; monocarps smooth; seeds ca. 17.5 mm long | G. latestigma C.E.C.Fisch. |
24b | Young branches (very) sparsely hairy; leaf laminas 10.5–13.5(–16) cm long, 3–5 cm wide; monocarps very finely rugulose; seeds ca. 9.5 mm long | G. repevensis Pierre ex Finet & Gagnep. |
23b | Monocarps red | 25 |
25a | Fruits without persistent calyx; seeds subglabrous to hairy | G. tapis Miq. |
25b | Fruits with persistent calyx; seeds glabrous | 26 |
26a | Leaf laminas 1.5–4 cm wide, ± glossy adaxially | G. elegans Ast |
26b | Leaf laminas 3.5–8.5 cm wide, (±) matt adaxially | 27 |
27a | Monocarps glabrous; monocarp stipes 4.5–13 mm long | G. tamirensis Pierre ex Finet & Gagnep. |
27b | Monocarps very sparsely hairy; monocarp stipes 2–3 mm long | G. rotundisepalus M.R.Hend. |
The authors would like to thank Mr Abdulromea Baka and Mr Isma-ael Sama-ae for field assistance, Ms Arisa Somphrom for the line drawings, and Dr Thomas Couvreur, Prof. David Johnson and two anonymous reviewers for their invaluable input. The first author would like to thank the Faculty of Science, Prince of Songkla University, for financial support.