Malesian Euphorbiaceae Descriptions

1. ACTEPHILA (Phyllanthaceae)

M. Heijkoop & P.C. van Welzen

Heijkoop, M. & P.C. van Welzen. 2017. A revision of the genus Actephila (Phyllanthaceae) in the Malesian Region. Blumea 62: 7–25.

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Genus description

Key to the species

Species descriptions

Excluded species

Actephila L.

    Actephila Blume, Bijdr. 12 (1826) 581; Endl., Gen. Pl. 16 (1840) 1123; Miq., Fl. Ned. Ind. 1, 2 (1859) 356; Müll.Arg., Linnaea 32 (1863) 77; in DC., Prodr. 15, 2 (1866) 221; Benth., Fl. Austral. 6 (1873) 88; Kurz, Forest Fl. Burma 2 (1877) 340; Benth. in Benth. & Hook.f., Gen. Pl. 3 (1880) 269; Hook.f., Fl. Brit. India 5 (1887) 282; F.M.Bailey, Queensl. Fl. 5 (1902) 1413; J.J.Sm., Meded. Dept. Landb. Ned.-Indië 10 (1910) 45; Pax & K.Hoffm. in Engl., Pflanzenr. VI.147.xv (1922) 191; Ridl., Fl. Malay Penins. 3 (1924) 196; Gagnep. in Lecomte, Fl. Indo-Chine 5 (1927) 530; Backer & Bakh.f., Fl. Java 1 (1963) 470; Airy Shaw, Kew Bull. 25 (1971) 496; Kew Bull. 26 (1972) 208; Whitmore, Tree Fl. Malaya 2 (1973) 51; Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 25; Kew Bull., Addit. Ser. 8 (1980a) 21; Muelleria 4 (1980b) 217; Kew Bull. 35 (1980c) 586; Kew Bull. 36 (1981) 248; Kew Bull. 37 (1982) 4; Alphabet. Enum. Euphorb. Philipp. Isl. (1983) 3; G.L.Webster, Ann. Missouri Bot. Gard. 81 (1994) 37; Govaerts, Frodin & Radcl.-Sm., World Checkl. Bibliogr. Euphorb. 1 (2000) 112; Radcl.-Sm., Gen. Euphorbiacearum (2001) 10; Chayam. in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 34; P.I.Forst., Austrobaileya 7 (2005) 63; Voronts. & Petra Hoffm., Kew Bull. 63 (2008) 46; G.L.Webster in Kubitzki, Fam. & Gen. Vasc. Pl. 11 (2014) 66; Heijkoop & Welzen, Blumea 62 (2017) 9. — Savia Willd. sect. Actephila (Blume) Baill., Étude Euphorb. (1858) 571, t. 26, f. 24. — Type (see note 1): Actephila javanica Miq. (see note 1) [= Actephila excelsa (Dalzell) Müll.Arg.].

    Lithoxylon Endl., Gen. Pl. 16 (1840) 1122; Baill., Étude Euphorb. (1858) 590; Müll.Arg. in DC., Prodr. 15, 2 (1866) 232. — Type: Lithoxylon lindleyi Steud. [= Actephila lindleyi (Steud.) Airy Shaw].

    Anomospermum Dalzell, Hooker’s J. Bot. Kew Gard. Misc. 3 (Aug. 1851) 228, nom. illeg. (non Miers, Jan. 1851, Menispermaceae). — Type: Anomospermum excelsum Dalzell [= Actephila excelsa (Dalzell) Müll.Arg.].

(Sub)shrubs to trees, monoecious; long internodes often alternating with several short ones, flowering branches round in section, with scars of fallen leaves, often flattened apically, striate when dry. Indumentum of short, simple, straight hairs, generally most parts subglabrous or glabrescent. Stipules stiff, apex acute, early caducous. Leaves alternate to often subopposite near branch ends, simple; petiole round but flat or channelled above, completely pulvinate or basally and apically pulvinate; blade elliptic, symmetric, papyraceous to pergamentaceous, margin entire, recurved, adaxially glabrous; venation raised on both sides when dry, pinnate, secondary nerves looped and joined near the margin, tertiairy veins and veinlets distinct, reticulate. Flowers solitary, paired or fascicled, generally axillary, when cauli- or ramiflorous then often in short racemes on thickened brachyblasts, 5-merous; sepals 5, imbricate, basally united, margin entire, outside somewhat sericeous, inside generally glabrous; petals 5 (Malesia), much smaller than sepals, obovate or obtriangular, base narrow, glabrous on both sides; disc a thin or fleshy ring around stamens or ovary, somewhat lobed. Staminate flowers: pedicel short; stamens 5, free or connate at base, anthers dorsibasifixed, 2-thecate, thecae parallel, opening latrorse via lengthwise slits; pistillode a dot of hairs or 3-branched and glabrous. Pistillate flowers: sepals persistent; petals semi-persistent; ovary 3-locular, ellipsoid, 2 ovules per locule, style absent, stigmas short, 3-branched, apically split, tips slightly thickened, bent horizontally, persistent. Fruits capsules, subglobose, generally apically flattened, 3-lobed, opening completely septicidally and loculicidally; pedicels round in cross section; sepals in some species enlarging; petals often caducous; wall usually knobbly, with 6 generally distinct lines of dehiscence, outer layer detaching from inner layer, latter somewhat woody; columella T-shaped, basally thickened. Seeds abaxially round, side walls straight, trigonous, smooth, without aril or caruncle; sometimes with widened abnormally shaped seeds when 1 ovule develops per locule, adaxially round with a longitudinal rib dividing it in 2 equal parts, abaxially flat, widely V-shaped, somewhat diamond shaped in cross section.

    Distribution — An Asian to Australasian and West Pacific genus of c. 25 species. In Malesia eleven species occur.

    Note — Blume (1826) only published the genus name, he never described a species However, he noted that the specimen he described came from the island Nusa Kambangan near Java. The only species Miquel described from this island, with reference to Blume, is A. javanica Miq. (basionym Savia actephila Hassk.) and accordingly, we selected a specimen of Blume from Nusa Kambangan as lectotype of Savia actephila and as type of the genus Actephila.

Key to the species

1. Actephila alanbakeri Welzen & Ent

    Actephila alanbakeri Welzen & Ent in Ent et al., Bot. Stud. 57, 6 (2016) 2; Heijkoop & Welzen, Blumea 62 (2017) 11, fig. 1, map 1. — Type: SNP (van der Ent et al.) 38539 (holo SNP; iso L), Malaysia, Sabah, Kinabalu Park, Nalumad. Paratype: SAN (Sugau & Seligi) 145750 (L, SAN), Malaysia, Saba, Kudat Distr., Malawali Island.

Shrubs to trees, up to 3 m high, d.b.h. up to 6 cm; flowering branches angular when dry, glabrous, 2–8 mm diam. Indumentum on brachyblasts and sepals, otherwise glabrous. Stipules triangular to deltoid or broadly ovate, 1.1–1.6 by (0.5–)0.8–1.3 mm. Leaves: petioles 4–30 mm long, green to maroon, heavily wrinkled when dry, pulvinate on both sides to completely pulvinate, pulvini becoming corky with age; blade narrowly ovate to narrowly elliptic, 8–24.5 by 1.6–4.9(–7.2) cm, 3.3–5.9 times as long as wide, papyraceous to pergamentaceous, base acute to obtuse, margin flat, apex acute or gradually tapering into a widely cuspidate apex, tip acute to obtuse, adaxially green, abaxially pale green, glabrous; nerves 15–20(–26) per side. Flowers solitary, on small brachyblasts with age; sepals apex acute to obtuse; disc a narrow thin ring. Staminate flowers c. 3 mm diam; pedicel short, not widening towards apex, glabrous; sepals ovate, widest just below the middle, reddish; petals apex truncate to somewhat wavy, glabrous, white to pinkish; stamens connate at base. Pistillate flowers (from photograph): sepals dark pink, ovate to broadly ovate, widest at base, c. 1 by 1.4 mm (as observed in fruit); petals white; disc yellow; ovary glabrous, stigmas yellow. Fruits green (immature) to brown (mature), 1.4–2 cm diam, pendent, glabrous; pedicels 7–33 by 0.75–1.5 mm, usually not widening towards apex; sepals not elongating; fruit wall with corky wrinkled surface, 3-lobed, lines of dehiscence consisting of 6 deep wide sutures, distinct, outer layer with strongly reticulate lines on the inside, somewhat elevated, somewhat woody, thick; columella c. 7 mm long, somewhat fibrous after dehiscence, basally completely covering disc and hanging over the base of sepals. Seeds c. 10 mm long, c. 8 mm wide, c. 7 mm high; widened seeds not seen.

    Distribution — Borneo and Philippines.

    Habitat & Ecology — Ultramafic forests, mangrove forests, along the beach, on gentle slopes. Soil: Ultramafics or limestone. Altitude: sea-level to 90m. Fruiting: June, August, October.

    Notes — 1. Typical for this species are the fruits with a wrinkled, thick, somewhat corky fruit wall when dry, and 6 wide sutures when fresh. The base of the columella in adult fruits is relatively wide and completely covering the persistent disc, the calyx is persistent but indistinct with small sepals. Additionally these plants are able to survive on ultramafic soils.

2. This species is hard to distinguish from A. lindleyi vegetatively, however, the stipules are in general shorter and narrower, the leaf blades have an acute or obtuse base instead of an attenuate base, and the species has in general 15–20 nerves per side of the midrib, while A. lindleyi only has in general 7–14 nerves per side of the midrib. Actephila alanbakeri is found in Borneo and in the Philippines, while A. lindleyi is known from New Guinea and the Moluccas.

3. The species was only known from ultrabasic soil in Sabah. Recently, two specimens from the Philippines (PNH (Celestino & Castro) 1932 from Mindoro, and PPI (Madulid & Majaducon) 36047 from Danjugan Island) could be added. However, it is unclear if these specimens were also growing on ultrabasic soil. Mindoro has ultrabasic soil, but for Danjugan information is lacking. The PNH specimen has no ecological information, and the PPI label is confusing, it indicates that the specimen was collected in beech and mangrove forest, but at an altitude of 50 m.

2. Actephila discoidea Heijkoop & Welzen

    Actephila discoidea Heijkoop & Welzen, Blumea 62 (2017) 11, fig. 2, map 2. — Type: Takeuchi & Towati 14639 (holo L; iso probably in A, LAE), Papua New Guinea, Morobe Province, Kamiali Wildlife Management Area, ridge inland of Cape Roon, slopes above side baret of the Ariwiri River, near 07º19.6’S, 147º07.5’E.

Shrubs to tree(let)s, 1–5 m high, monocaulous to sparsely branched; flowering branches 2–4 mm diam, with brown lenticels, sometimes somewhat puberulous at branch tips. Indumentum: most parts subglabrous. Stipules deltoid to broadly ovate, 1.5–3 by 1.2–2 mm, glabrous. Leaves: petioles 0.5–3.8 cm long, pulvinate at both ends, often glabrous, sometimes slightly sericeous, striate when dry, sometimes with transverse grooves, somewhat corky when older; blade ovate to elliptic, 11–30.5 by 2.6–9.3 cm, 2.6–4.3(–5.1) times as long as wide, fleshy when fresh, papyraceous when dry, base obtuse to acute, margin flat, apex gradually tapering, widely caudate to cuspidate, tip acute, adaxially dull to glossy, light to dark green, abaxially paler green than adaxially to yellow-green or slightly glaucous, glabrous; nerves 9–14 per side Flowers solitary, on brachyblasts when older. Staminate flowers not seen. Pistillate flowers not seen, observations from fruit: sepals broadly ovate, widest at base, c. 3 by 2–2.5 mm, apex acute to obtuse, often glabrous on both sides; disc a thick fleshy, wide ring, substantially covering the base of sepals in fruit. Fruits dull, immature glaucous green to green, 1.5–1.7 cm diam, erect, glabrous; pedicels 7–20(–25) by 1–2 mm, usually not thickening towards apex, straight, edges smooth, glabrous (to slightly sericeous); sepals not elongating, reflexed; wall with knobbly surface, thin, lines of dehiscence not elevated; columella 6–11 mm long, fibrous after dehiscence. Seeds 8–12 mm long, 5–9 mm wide, 5–7 mm high; widened seeds not seen.

    Distribution — New Guinea.

A. discoidea: squares; A. dolichopoda: triangles; A. emarginata: dots.

    Habitat & Ecology — Primary lowland (dipterocarp) rain forests. Soil: clay, rocks, ultrabasic soil. Altitude: sea-level to 430 m. Fruiting in March, April, July, August, November.

    Notes — Typical for the species are the leaves with an obtuse to acute base and a gradually tapering long apex; fruits on short, straight, sturdy pedicels; a thick, fleshy disc, visible after fruit dehiscence.

3. Actephila dolichopoda Airy Shaw

    Actephila dolichopoda Airy Shaw, Kew Bull. 32 (1978) 380; Kew Bull., Addit. Ser. 8 (1980a) 22; Heijkoop & Welzen, Blumea 62 (2017) 13, fig. 3, map 2. — Type: Carr 11774 (holo K; iso A, BM, L), Papua New Guinea, Kanosia.

Shrubs to trees, up to 5 m high; flowering branches 1–1.5 mm diam, covered with brown lenticels and hair stubbles. Indumentum: subglabrous with very short, papillae-like hairs on especially the younger branches, petioles, leaf blades; sericeous hairs on fruiting sepals. Stipules deltoid or broadly ovate to triangular or ovate, 0.8–1.6 by 0.5–0.9 mm, glabrous. Leaves: petioles 5–27(–37) mm long, glabrous, pulvinate at both ends, pulvini becoming corky with age; blade elliptic to somewhat obovate, 7–16.3 by 2.7–6.8 cm, 2.4–2.9 times as long as wide, papyraceous, base acute, margin flat, apex acuminate to cuspidate, adaxially mid-green, abaxially paler mid-green, glabrous; nerves 10–11 per side. Flowers solitary. Staminate flowers not seen. Pistillate flowers not seen, observations from fruit: sepals obovate, widest just above the middle, c. 6 by 2.5 mm, apex obtuse, outside somewhat sericeous; petals damaged; disc a distinct, thick, fleshy, wide ring, substantially covering the base of the sepals when fruiting. Fruits c. 1.8 cm diam, green; pedicels 27–85 by 0.75–1(–2) mm, straight, often widening towards apex, glabrous; sepals not enlarging, reflexed, outside somewhat sericeous, glabrescent; wall with a very knobbly surface, c. 2 mm thick, glabrous; columella 9–13 mm long, basally completely covering disc. Seeds 12–13 mm long, 7–9.5 mm wide, 6–8 mm high, widened seeds c. 12 mm long, c. 11 mm wide, 6–7 mm high.

    Distribution — Papua New Guinea (endemic in the Central Prov.).

A. dolichopoda: triangles; A. discoidea: squares; A. emarginata: dots.

    Habitat & Ecology — Lowland rain forest to (slightly deciduous) hill forest; soil: volcanic substrate. Altitude: sea-level up to 10 m (and likely higher). Flowering: April; fruiting: April, July.

    Notes — Typical for this species are the fruits on long, straight pedicels of up to 8.5 cm length. After dehiscence a fleshy nectar disc and a long columella, of up to 13 mm long, are visible with the columella extremely widening at the base completely covering the large disc. Further the fruit wall is somewhat thicker and more knobbly compared to the other species. Also remarkable are the branches covered with short, stiff, papillae-like hairs, while other Actephila species in this region either have short simple hairs or are glabrous on the branches. The stipules are on average short and narrow.

4. Actephila emarginata Heijkoop & Welzen

    Actephila emarginata Heijkoop & Welzen, Blumea 62 (2017) 13, fig. 4, map 2. — Type: Brass 29350 (holo L), Papua New Guinea, Morobe District, Oomsis.

Shrubs, 1–2 m high, monocaulous or sparsely branched; flowering branches 2–4 mm diam, puberulous. Indumentum on most parts subglabrous. Stipules triangular to ovate, 1.4–1.7 by 1–1.2 mm, glabrous. Leaves: petioles 3–6 mm long, completely pulvinate, striate when dry, sometimes with corky edges, glabrous to densely puberulous; blade obovate, sometimes elliptic, 9.5–27 by 4.6–10 cm, 1.6–2.7 times as long as wide, papyraceous, base emarginate to sometimes more obtuse in a few leaves, margin flat, apex acuminate to cuspidate, adaxially dull, dark green, abaxially pale green, puberulous on midrib and venation (more so towards leaf base); nerves 8–13 per side. Flowers solitary, green. Staminate flowers not seen. Pistillate flowers not seen, data from fruit; sepals broadly ovate or ovoid, widest just below the middle to at base, 2–2.5 by 1.5–2 mm, apex obtuse; petals not seen; disc a wide fleshy ring, substantially covering sepal base in fruit. Fruits 1.5–2 cm diam, green; pedicels 25–50 by 0.5–1 mm, thickening towards apex, smooth to edges somewhat striate when dry; sepals not elongating, horizontal to reflexed; wall with knobbly surface, lines of dehiscence not elevated, thin, glabrous; columella c. 6 mm long, basally completely covering the disc. Seeds immature flat, half-moon-shaped, c. 8 mm long, 1–2 mm wide, c. 5 mm high, mature not seen.

    Distribution — New Guinea (only known from the type, endemic in area around Oomsis).

A. emarginata: dots; A. discoidea: squares; A. dolichopoda: triangles.

    Habitat & Ecology — Rain forest on flood bank of stream. Altitude: 100 m. Flowering: May; fruiting: May.

    Notes — Typical for this species is the leaf base, which is generally emarginate, which do not occur in any of the other Actephila species in this region. The petioles are short and often strongly puberulous and the fruits have a relatively long, straight pedicel of up to 5 cm length.

5. Actephila excelsa (Dalzell) Müll.Arg.

    Actephila excelsa (Dalzell) Müll.Arg., Linnaea 32 (1863) 78; in DC., Prodr: 15, 2 (1866) 222; Hook.f., Fl. Brit. India 5 (1887) 282; Pax & K.Hoffm. in Engl., Pflanzenr. VI.147.xv (1922) 191; Ridl., Fl. Malay Penins. 3 (1924) 196; Gagnep. in Lecomte, Fl. Indo-Chine 5 (1927) 535; M.R.Hend., J. Malayan Branch Roy. Asiat. Soc. 17 (1939) 68; Backer & Bakh.f., Fl. Java 1 (1963) 470; Airy Shaw, Kew Bull. 26 (1972) 209; Whitmore, Tree Fl. Malaya 2 (1973) 52; Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 25; Chayam. in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 35, fig. 5; Heijkoop & Welzen, Blumea 62 (2017) 14, map 3. — Anomospermum excelsum Dalzell, Hooker’s J. Bot. Kew Garden Misc. 3 (1851) 228. — Actephila excelsa (Dalzell) Müll.Arg. var. genuinum Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 192, nom. inval. — Actephila excelsa (Dalzell) Müll.Arg. var. excelsa: Airy Shaw, Kew Bull. 26 (1972) 209. — Type: Dalzell s.n. (holo K, barcode K000186757), India, Syhadrensi?

    [Croton laevigatum Wall., Numer. List (1828) 7749 (non 7735), nom. nud., non Vahl].

    Actephila neilgherrensis Wight, Icon. Pl. Ind. Orient. 5 (1852) 29, t. 1910. — Type: Wight KD 2574 (K), [India,] Neilgherries, Coonoor.

    Savia actephila Hassk., Cat. Hort. Bogor. (1844) 243; Baill., Étude Euphorb. (1858) 571. — Actephila javanica Miq., Fl. Ned. Ind. 1, 2 (1859) 356; Müll.Arg. in DC., Prodr. 15, 2 (1866) 222; J.J.Sm., Meded. Dept. Landb. Ned.-Indië 10 (1910) 46; Ridl., Fl. Malay Penins. 3 (1924) 196; Backer & Bakh.f., Fl. Java 1 (1963) 470; Airy Shaw, Kew Bull., Addit. Ser. 8 (1980a) 22; Kew Bull. 37 (1982) 4; Chayam. in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 38, fig. 6. — Actephila excelsa (Dalzell) Müll.Arg. var. javanica (Miq.) Pax & K.Hoffm. in Engl., Pflanzenr. VI.147.xv (1922) 192; Merr., Enum. Philipp. Fl. Pl. 2 (1923) 390; Pl. Elmer. Born. (1929) 139; Airy Shaw, Kew Bull. 26 (1972) 209; Whitmore, Tree Fl. Malaya 2 (1973) 52; Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 25; Kew Bull., Addit. Ser. 8 (1980a) 22; Kew Bull. 36 (1981) 248; Alphabet. Enum. Euphorb. Philipp. Isl. (1983) 3. — Lectotype (designated by Heijkoop & Welzen 2017): Blume 1670 (L), Java.

    Actephila bantamensis Miq., Fl. Ned. Ind. 1, 2 (1859) 356. — Syntypes: Hasskarl s.n. (perhaps U fragment), [Indonesia,] Java; Zollinger 1662 (G-DC, P, 2 sheets, fragment U?), [Indonesia,] Java, Zuid Bantam (S Bantam).

    Actephila major Müll.Arg., Linnaea 32 (1863) 77; in DC., Prodr. 15, 2 (1866) 221. — Type: Zollinger 512 (holo B, lost; iso perhaps fragment in G-DC as Zollinger s.n.), [Indonesia,] Java.

    [Savia zeylanica Baill., Étude Euphorb. (1858) 571, nom. nud. —] Actephila zeylanica Baill. ex Müll.Arg., Linnaea 32 (1863) 77; in DC., Prodr. 15, 2 (1866) 221. — Actephila excelsa (Dalzell) Müll.Arg. var. zeylanica (Baill. ex Müll.Arg.) Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 192. — Type: Thwaites 2532 (G-DC, P, 4 sheets), Ceylon (= Sri Lanka).

    Actephila thomsonii Müll.Arg., Linnaea 34 (1865) 65; in DC., Prodr. 15, 2 (1866) 222. — Actephila excelsa (Dalzell) Müll.Arg. var. thomsonii (Müll.Arg.) Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 192. — Type: G. Thomson s.n. (not located), India, Maisor & Carnatic.

    Actephila gigantifolia Koord., Meded. Lands Plantentuin 19 (1897) 579, 625 (description); Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 194. — Type: Koorders 16937 (holo L), [Indonesia,] Celebes, bivak Totok.

    Actephila minahassae Koord., Meded. Lands Plantentuin 19 (1897) 579, 625 (description); Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 195. — Type: Koorders s.n. (not found), [Indonesia,] Celebes, Ratahan.

    Pimeleodendron dispersum Elmer, Leafl. Philipp. Bot. 1 (1908) 308 (‘dispersa’); Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.v (1912) 56 (excluded species). — Actephila dispersa (Elmer) Merr., Philipp. J. Sci 4, Bot. (1909) 276. — Type: Elmer 7245 (holo PNH, lost; iso K, 2 sheets, L), Philippines, Leyte, Palo.

    Actephila gitingensis Elmer, Leafl. Philipp. Bot. 3 (1910) 903; Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 25; Alphabet. Enum. Euphorb. Philipp. Isl. (1983) 3. — Type: Elmer 12141 (holo PNH, lost; iso A, E, G, GH, HBG, K, L, MO, US), Philippines, Sibuyan Island, Cadiz Prov., Magallanes (Mt. Giting-giting).

    Actephila magnifolia Elmer, Leafl. Philipp. Bot. 3 (1910) 903; Merr., Enum. Philipp. Fl. Pl. 2 (1923) 390; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 193; Airy Shaw, Alphabet. Enum. Euphorb. Philipp. Isl. (1983) 3 — Type: Elmer 12303 (holo PNH, lost; iso A, E, G, HBG, K, L, MO, US), Philippines, Sibuyan Island, Cadiz Prov., Magallanes (Mt. Giting-giting).

    Actephila dolichantha Croizat, J. Arnold Arbor. 23 (1942) 30. — Type: C.W. Wang 79253 (holo A, not seen), China, Yunnan, Kuen-ger, Che-li Hsien (for inclusion under A. excelsa see Govaerts et al. 2000).

   Actephila excelsa (Dalzell) Müll.Arg. var. brevifolia N.Balach., Mahesw. & Chakrab., J. Econ. Taxon. Bot. 33 (2009) 717, fig. 1. — Type: B. Maheswaran 990 A (holo AURO, not seen; iso AURO, not seen), India, Andamans, North Andaman Island, Saddle Peak. See note 3.

Shrubs to tree(let)s, up to 10(–15) m high, d.b.h. up to 30 cm; flowering branches 2–9(–20 when rami- or cauliflorous) mm diam, with (light) brown lenticels, glabrous to puberulous, more puberulous towards apex. Outer bark pale tan to greyish to greenish yellow to reddish, smooth to with fine vertical fissures to scaly; inner bark green to khaki-green to yellow; sapwood white to pale orange. Indumentum: most parts subglabrous to somewhat hairy. Stipules triangular or ovate to deltoid or broadly ovate, (1–)1.5–3(–3.5) by 1–2 mm, slightly sericeous outside to often glabrous. Leaves alternate to subopposite at branch ends; petioles 5–95 mm long, glabrous to slightly to completely puberulous, brown, pulvinate at both ends, pulvini becoming corky with age; blade elliptic (to ± obovate), (4-)5.5–35.5 by (1.1–)1.9–13.5(–15.9) cm, 1.9–3.6(–8) times as long as wide, papyraceous to pergamentaceous, base acute to obtuse, margin flat, apex cuspidate (to acuminate or rarely acute), adaxially glossy, light to dark green, abaxially paler green than adaxially, sometimes somewhat puberulous on midrib and secondary venation, more so towards base; nerves (5-)7–16 per side. Flowers solitary to fascicled, staminate ones often cauliflorous or ramiflorous, pistillate ones axillary, on brachyblasts after several seasons (up to 2 cm diam on thick branches); pedicels green, widening towards apex, glabrous to sericeous; sepals light green to white (with pink base), margin sometimes slightly membranous, apex acute to obtuse; petals: apex truncate to obtuse or rounded, erose, white to greenish white; disc a narrow thin ring, light yellow. Staminate flowers 2–3 mm diam; pedicels c. 2 mm long; sepals ovate to elliptic to diamond shaped, widest just below to in the middle, 1.3–2 by 1.2–2 mm, green to light green; petals 1–2 by 0.8–1 mm; stamens free, greenish to white, anthers (light) yellow. Pistillate flowers 3–5 mm diam, white to greenish; pedicel 8–31(–53) mm long; sepals ovate to elliptic to somewhat diamond shaped to obovate, widest just below to just above the middle, (1.5–)2–4(–5) by (1–)1.5–3(–3.5) mm, green to greenish white; petals (0.6–)0.7–2 by (0.3–)0.5–1.3 mm, sometimes somewhat split into two; disc slightly covering base of sepals in fruit; ovary 0.8–2.5 by 0.9–3 mm, green to green-white, glabrous, stigmas 0.4–0.6 mm long, basally pinkish, apically light green. Fruits 1.3–2 cm diam, pendent, green (immature) to brown (mature); pedicels elongating up to 75 mm long, (0.4–)0.7–1.5 mm diam, often curved, glabrous (to slightly sericeous), somewhat striate when dry; sepals not elongating, reflexed to flat when dry, dark green, outside sometimes slightly sericeous, often glabrous; fruit wall somewhat knobbly, lines of dehiscence sometimes slightly elevated, thin, glabrous (to slightly sericeous); columella 5–8 mm long, basally somewhat thickened but not completely covering disc and base of sepals, fibrous after dehiscence. Seeds 7–12 mm long, 5–10 mm wide, 5–9 mm high, widened seeds (7–)8–13 mm long, (6.5–)9–11(–14) mm wide, 5–8(–9) mm high.

    Distribution — India, Sri Lanka, Myanmar, S. China, Thailand, Indochina, Malay Penisula, Sumatra, Java, Borneo, Philippines, Sulawesi.

    Habitat & Ecology — Primary mixed lowland dipterocarp rain forest, evergreen forest, transition to deciduous forest, secondary forest, hill forest, kerangas forest, along rocky seashores, rivers and roads. Soil: limestone, loam, sandy soil, ultrabasic, sandy clay, granite-derived, basalt. Altitude: sea-level to 500(–1350) m. Flowering and fruiting: the whole year through.

    Vernacular names — Malay Peninsula: Kelapa Tupai. Sumatra: Baner etem, Pehailigei, Toetoeg getah. Philippines: Manikusan.

    Notes — 1. A highly variable species, which greatly varies in leaf size and petiole length. Leaf blade shapes vary from elliptic to somewhat ovate or obovate, apices vary from acute to cuspidate. Fruiting pedicels are usually around 2–3 cm long, curved and widening towards apex, but longer pedicels are observed losing their curved shape. The indumentum varies from almost completely absent to being heavily puberulous on most parts. Various groups of specimens have somewhat distinct characters:

    a. The northern latitudes (N & SW Thailand) and the specimens in the western part of the distribution (India, Sri Lanka) have generally smaller leaves and short petioles (formerly var. excelsa; Chayamarit 2005), while in most of Malesia, but also S Thailand, the petioles are long(er) and the leaf blades can be quite large (see also under f; formerly var. javanica; Chayamarit 2005), but small blades are also found throughout the distribution (see under b).

    b. Forbes 3112a, Korthals s.n., SAN (Sugau & Diwol) 141043, SAN( Maikin, Donggop & Francis) 130671, BRUN (Salleh Ismail) 16862, PPI (Stone, Reynoso & Fernando) 6964, de Vogel 2771, University of San Carlos 98 have small leaf blades, 5.5–14.5 by 1.9–5.8 cm and short petioles, 0.4–2.7 cm long. The samples are found in the eastern part of the species distribution: Sumatra, Java, Borneo and the Philippines. PPI 6964 (Philippines) is also special because of the somewhat elevated reticulate lines on the outside of the fruit wall, instead of the usual knobbly surface.

    c. KEP FRI (Loh) 19212 (Malay Peninsula) has very narrow leaves, up to 8 times longer than wide, also the fruits seem slightly smaller and the seeds are below average size. The plant is growing near a waterfall, which may cause the slightly different appearance.

    d. Koorders 26891 and 30269 have long fruiting pedicels, 6–7.5 cm long, 4–7 cm in Koorders 39682, while Koorders 2738 has flowering pedicels of c. 5.3 cm long. All specimens are from Java. No other specimens have fruiting pedicels exceeding 5.5 cm. The petioles are relatively short for the species with a general length of 7–28 mm. However, the leaf blade sizes of 10–25 by 4.5–9 cm typically fit in with the rest of the specimens.

    e. Afriastini 1243, Rappard 168, and Backer 36980, all from Java, show somewhat smaller stipules, petioles and leaf blades: Stipules sizes vary between 0.8–1.2 by 1–1.2 mm, petiole length 2–18 mm long, and leaf blade sizes of 4–14.5 by 2–5.5 cm, which are 2–2.7 times as long as wide. Apices vary between acute and acuminate, never extending up to cuspidate.

    f. The types of A. gigantifolia, Koorders 16937 (Sulawesi), and A. magnifolia, Elmer 12303 (Philippines), show enormous leaf blades, up to 35 by 17 cm, much larger than all other specimens of Actephila. However, the specimens still show the typical bend in the upper pulvinus.

2. Actephila puberula Kurz, from the Andaman Islands, has often been treated as synonym of A. excelsa, but the leaf base is emarginate to rounded instead of obtuse to actue and the ovary is densely hairy instead of glabrous. Therefore, A. puberula (with A. excelsa var. puberula (Kurz) Pax & K.Hoffm. as synonym) is here considered to be a distinct species. The fruits of A. puberula can be up to 2.5 cm diam (instead of up to 2 cm) and the surface is more knobbly than in A. excelsa.

3. Actephila excelsa var. brevifolia is added to the synonymy, because it has the typical leaf base of A. excelsa and does not really differ the description above other then sometimes slightly smaller leaf blades (with consequently slightly fewer secondary nerves).

6. Actephila latifolia Benth.

    Actephila latifolia Benth., Fl. Austral. 6 (1873) 89; F.M.Bailey, Queensl. Fl. 5 (1902) 1414; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 194; Airy Shaw, Kew Bull. 35 (1980c) 587; Kew Bull. 37 (1982) 4; P.I.Forst., Austrobaileya 7 (2005) 74, Fig. 5; Heijkoop & Welzen, Blumea 62 (2017) 17, map 4. — Lectotype (designated by Airy Shaw 1980c): Daemel s.n. (holo K, photo BRI), Australia, Queensland, Cape York; other former syntype: Dallachy s.n. (not seen), [Australia,] Queensland, Rockingham Bay.

Shrubs to tree(let)s, up to 5 m high, d.b.h. up to 5 cm; flowering branches 1–3 mm diam, with few brown lenticels, somewhat sericeous at tips. Bark nondescript; blaze thick, pinkish-red; wood dense, cream-yellow. Indumentum: subglabrous on branches, brachyblasts and sepals. Stipules deltoid to broadly ovate, 0.8–1.3 by 0.8–1.3 mm, glabrous, often somewhat folded horizontally. Leaves alternate to sub-opposite at branch ends; petioles 3–22 mm long, glabrous, completely pulvinate to pulvinate at both ends; blade 6.5–14.9 by 3.2–6.9 cm, 1.9–2.4(–3) times as long as wide, pergamentaceous, base slightly attenuate to acute, margin flat, apex obtuse to acute to slightly acuminate, abaxially glabrous; nerves 7–10 per side. Flowers white to cream-yellow, solitary to paired or fascicled, on older branches on small brachyblasts; pedicels not widening towards apex, glabrous; sepals: apex rounded or somewhat obtuse; petals: apex rounded, not erose; disc a thick, fleshy, wide ring. Staminate flowers c. 4 mm diam; pedicels c. 7 mm long; sepals ovate to elliptic, c. 2.8 by 1.8 mm, margin slightly membranous; petals c. 1 by 0.5 mm; disc hanging over the base of the sepals; stamens: filaments c. 2 mm long. Pistillate flowers c. 4 mm diam; pedicel 6–35 mm long; sepals ovate to elliptic to obovate, widest just below to in the middle, 2–3.3 by 1–3.3 mm; petals c. 1.2 by 0.6 mm; disc pale yellow, substantially covering base of sepals in fruit; ovary c. 0.8 by 1–1.4 mm, glabrous, stigmas 0.8–2 mm long. Fruits c. 10 mm diam, green; pedicels 15–35 by 0.7-0.8 mm, barely widening towards apex, glabrous, smooth to striate when dry; sepals slightly enlarging, 4.5–5(–5.5) by 2.5–4.5(–5) mm, glabrous on both sides, reflexed; wall with knobbly surface, thin, glabrous; columella 6-8 mm long. Seeds c. 8.5 mm long, c. 7 mm wide, c. 7 mm high; widened seeds not seen.

    Distribution — Moluccas (Aru Isl.), Lesser Sunda Islands (Tanimbar, Timor), Australia (Queensland).

A. latifolia: triangles; A. lindleyi: dots.

    Habitat & Ecology — Riverine to rainforest bordering savannah, semi-deciduous to complex notophyll vine-forest. Soil: sand, granite. Altitude: 10–480 m. Flowering: January; fruiting: March–May, October.

    Vernacular name — Aru Islands (P. Trangan): Balabal.

    Notes — Typical for this species are the small deltoid stipules, which often have a horizontal fold in the middle. The petioles are short, the leaf blades are relatively constant in size and generally up to 15 by 7 cm with an acute or obtuse apex. The calyx slightly enlarges in fruit to generally c. 5 by 4 mm. The persistent disc is wide, thick and fleshy, covering the base of the sepals. The septa often look weak and thin, not as sturdy as in other species, however, this may be due to underdevelopment.

7. Actephila lindleyi (Steud.) Airy Shaw

    Actephila lindleyi (Steud.) Airy Shaw, Kew Bull. 25 (1971) 496; Kew Bull., Addit. Ser. 8 (1980a) 23; Kew Bull. 35 (1980c) 587; Kew Bull. 37 (1982) 4; N.C.W.Beadle, O.D.Evans & Carolin, Fl. Sydney Reg., 3^rd ed. (1982) 351; P.I.Forst., Austrobaileya 7 (2005) 75, fig. 6; ; Heijkoop & Welzen, Blumea 62 (2017) 17, fig. 5, map 4. — Securinega nitida W.T.Aiton [, Hort. Kew. ed. 2, 5 (1813) 383, nom. nud.] ex Lindl., Coll. Bot. (1821) t.9 (non Willd.; see A.Juss., Euphorb. Gen. (1824) 14, in obs.). — Lithoxylon lindleyi Steud., Nomencl. Bot. ed 2, 2 (1841) 57, pro nom. nov. — Lithoxylon nitidum (W.T.Aiton ex Lindl.) Baill., Étude Euphorb. (1858) 590, nom. illeg., superfl.; Müll.Arg. in DC., Prodr. 15, 2 (1866) 232; Nadeaud, Énum. Pl. Tahiti (1873) 72. — Actephila nitida (W.T.Aiton ex Lindl.) Benth. & Hook.f. ex Drake, Ill. Fl. Ins. Pacif. (1892) 286, nom. illeg..; Fl. Polynésie Franç. (1893) 177; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 194. — Type: cult. in Hort. Kew., 1813-1821 (holo K, photo BRI), native of Otaheite [= Tahiti].

    Lithoxylon grandifolium Müll.Arg., Linnaea 34 (1865) 65; in DC., Prodr. 15, 2 (1866) 232. — Actephila grandifolia (Müll.Arg.) Baill., Adansonia 6 (1866) 330, 360, t. 10; Benth., Fl. Austr. 6 (1873) 89; F.M.Bailey, Queensl. Fl. 5 (1902) 1413; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xv (1922) 193; R.H.Anderson, Trees N.S.W., ed. 4 (1968), 218, 377; N.C.W.Beadle, O.D.Evans & Carolin, Fl. Sydney Reg., rev. ed. (1972) 252; P.I.Forst., Austrobaileya 7 (2005) 71, Fig. 4. — Type: F. Mueller s.n. (holo G-DC, BRI photo), [Australia, New South Wales,] Clarence River.

(Woody) herbs to shrubs to small trees, up to 9 m high, at least up to 10 cm diam; flowering branches 2–5 mm diam, glabrous to slightly sericeous at tip. Outer bark grey-brown to brown, smooth, with many, large, brownish, often somewhat raised lenticels to shallowly longitudinally fissured; blaze white; inner bark reddish brown, no exudate; wood cream to pale orange. Indumentum: most parts subglabrous. Stipules deltoid or broadly ovate to triangular or ovate, 1.2–3(–4) by 1–2 mm, straight, glabrous. Leaves alternate to almost opposite at end of branches; petioles 0.1–2.7 cm long, completely pulvinate, apex sometimes gradually tapering into base of blade, often with corky edges when older, often glabrous, sometimes slightly sericeous; blade elliptic to narrowly elliptic to sometimes somewhat obovate, 7.5–25(–30) by (2–)2.5–11cm, 1.7–4.8(–6.7) times as long as wide, papyraceous to pergamentaceous, base usually attenuate to rarely slightly more acute, margin flat, apex acute or somewhat obtuse to acuminate or cuspidate, adaxially dull to glossy green to dark green, abaxially lighter green than above, glabrous; nerves 7–14(–18) per side. Flowers often solitary to paired or fascicled, on brachyblasts when older, green-white to white; pedicels not widening towards apex, often glabrous, sometimes slightly sericeous; sepals: apex acute to obtuse; petals: apex truncate to rounded, erose; disc a thin narrow papery ring. Staminate flowers c. 3 mm diam, green to white with purple tinge; pedicel 3–4 mm long; sepals ovate, widest just below the middle, c. 2 by 1.5 mm, margin membranous; petals c. 1 by 0.5 mm; disc folded upwards at the edges; stamens: filaments c. 1.3 mm long, connate at base; small pistillode on staminal tube. Pistillate flowers c. 4 mm diam, white; pedicel c. 3 mm long; sepals broadly ovate or deltoid to ovate to elliptic, 1.3–2.5(–3) by 1.3–2(–2.5) mm, widest at base to in the middle; petals c. 0.7–1 by 0.25–0.5 mm; disc slightly covering the base of sepals in fruit; ovary c. 1 by 0.8–1.5 mm, glabrous, stigmas 0.4–0.6 mm long. Fruits immature globose, mature subglobose, 1–2 cm diam, light green (immature) to brownish green to red; pedicels 6–15(–40) by (0.5–)0.9–2(–2.5) mm, straight, sometimes with transverse small grooves, edges smooth to somewhat striate when dry, sometimes somewhat corky, glabrous; sepals not enlarging, completely glabrous, reflexed; wall with knobbly surface, thin, glabrous, lines of dehiscence not elevated; columella 5–8 mm long, fibrous after dehiscence. Seeds 7.5–10 mm long, (5–)6–8 mm wide, 4.5–6.5 mm high, widened seeds 7–8 mm long, 7–8 mm wide, 4–5 mm high.

    Distribution — Moluccas, New Guinea (incl. Aru Islands and New Britain), Solomon Islands (Santa Ysabel).

A. lindleyi: dots; A. latifolia: triangles.

    Habitat & Ecology — Primary lowland rain forest, riverine forest, Castanopsis-oak forest, old secondary forest, along rivers and sago swamps. Soil: (sandy) clay to limestone or coral sand. Altitude: sea-level to 580(–1370) m. Flowering: February–August, November; fruiting: January–October.

    Vernacular names — Aru Islands (Pulau Kobroor): Ay, Ay kway ngom. New Guinea: Papua: Krokaan (Maibrat); New Britain: Ugunape. Solomon Islands (Santa Ysabel): Susui.

    Notes — 1. The leaves in this species are highly variable, they range from short to long, narrow to wide, apices range between acute and cuspidate. However, the attenuate leaf blade base is constant (almost unique to this species, with the exceptions of A. ovalis and A. latifolia). Typical are the short petioles and short and somewhat sturdy, straight fruiting pedicels.

Exceptional forms are:

- Specimens from Indonesian Papua tend to have narrower leaves, Van Royen 5296 being narrowest, with leaf blades up to 6.7 times as long as wide. Also exceptional are the subterminal fascicles of relatively small fruits. Aet & Idjan 342 (1939) also has a very narrow base which gradually merges into the petiole, and the apex is gradually tapering into a widely cuspidate apex (latter also in BW (van der Sijde) 4161). Brass 32390 was collected at the exceptionally high altitude of 1370 m; it only deviates from other specimens in the leaf blades with an obtuse to somewhat acute apex.

- Several specimens have long petioles and/or pedicels. NGF (Frodin) 26551 has long pedicels (c. 4 cm long), but also somewhat longer petioles (up to 2 cm long). LAE (Wiakabu et al.) 50086 deviates in longer petioles (12–27 mm long), longer fruiting pedicels (c. 23 mm long) and an obtuse to acute leaf blade base (instead of attenuate). NGF (Katik) 46983 also has much longer petioles (25–48 mm long) and longer fruiting pedicels (c. 26 mm long), which are somewhat widening towards the apex. Also exceptional is the presence of indumentum (other A. lindleyi are subglabrous), especially the moderate indumentum on the petioles is unique.

2. Actephila mooreana Baill. is to be recognised as an accepted species and not a synonym of A. lindleyi. The latter species does not occur in Australia and differs in the characters leaf apex, leaf size, leaf margin, size of the disc, and pistilode. Leaf blades of A. mooreana are up to 17 cm long (often below 10 cm), with an acute to rounded apex and a wavy margin. The pistillode in the staminate flowers of A. mooreana is hairy. The disc in A. mooreana is wide and fleshy in staminate and pistillate flowers, pushing the sepals backwards in fruit. Fruits of A. mooreana are up to 1.3 cm diam. Actephila mooriana, the orignal spelling by Baillon (1866) was incorrect.

8. Actephila ovalis (Ridl.) Gage

    Actephila ovalis (Ridl.) Gage, Rec. Bot. Surv. India 9 (1922) 219; Ridl., Fl. Malay Penins. 3 (1924) 196; Gage, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 75 (1936) 519; M.R.Hend., J. Malayan Branch Roy. Asiat. Soc. 17 (1939) 68; Smitinand, Nat. Hist. Bull. Siam Soc. 20 (1962) 139; Airy Shaw, Kew Bull. 26 (1972) 210; Whitmore, Tree Fl. Malaya 2 (1973) 51; Chayam. in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 38, fig. 7, plate II: 1; Heijkoop & Welzen, Blumea 62 (2017) 19, fig. 6, map 5. — Dimorphocalyx ovalis Ridl., J. Straits Branch Roy. Asiat. Soc. 59 (1911) 178; Smitinand, Nat. Hist. Bull. Siam Soc. 20 (1961) 52, photo. — Syntypes: Curtis s.n. (SING), Malaysia, Lankawi, Kwah; Ridley 14890 (K, 2 sheets, SING), Malaysia, Perlis woods at Bukit Telor Jambu, Kanga.

(Sub)shrubs to small trees, up to 5 m high, stem brown; flowering branches 1–4 mm diam, with brown to whitish lenticels, puberulous. Indumentum of simple hairs, most parts subglabrous. Stipules narrowly ovate to narrowly triangular, 1.5–4 by 0.8–1.5 mm, outside strongly sericious when young, often glabrous. Leaves alternate to subopposite; petioles 2–6(–13) mm long, completely pulvinate, strongly puberulous; blade elliptic, 6.4–15.4 by 2.1–4.9 cm, 2.7–3.8(–4.4) times as long as wide, papyraceous, base attenuate to acute, margin wavy, apex narrowly to widely cuspidate, sometimes more acuminate, adaxially mid to dark green, abaxially slightly lighter green, sometimes somewhat puberulous basally on midrib; nerves 10–17 per side. Flowers solitary, on older branches on small brachyblasts; petals: apex truncate to rounded, erose; disc a thick fleshy ring. Staminate flowers c. 5 mm diam; pedicel 2–4 mm long, widening towards apex, glabrous to sericeous, cream-coloured; sepals ovate, widest just below the middle, c. 2 by 1.2–1.5(–2) mm, light greenish to cream, margin slightly membranous, apex rounded or somewhat obtuse, outside hairy; petals c. 2 by 0.9 mm, white; disc covering base of sepals, whitish; anthers yellow to orangish; pistillode small. Pistillate flowers green; sepals: lobes obovate, 2 smaller, c. 3.5 by 3 mm, 3 lagerall obovate, c. 5 by 4.5 mm, enlarging in fruit, green; petals 1.4–1.5 by 0.5–0.7 mm; disc substantially covering sepal base in fruit; ovary ovoid, slightly 3-lobed, c. 1.2 by 1.6 mm, greenish; style c. 0.5 mm long; stigmas c. 1.5 mm long, cream to purple, apically slightly split. Fruits 1.6–1.8 cm diam, (green to) pale green to yellowish green; pedicels (25–)30–70 by 0.4–0.9 mm, straight, not widening towards apex, often somewhat striate when dry, aprically remaining somewhat puberulous; sepals enlarged, elliptic, widest in the middle, (10–)13–22 by 9–15 mm, margin entire, apex rounded, venation parallel, 4–8 main veins, basally slightly hairy outside, otherwise glabrous; wall with knobbly surface, sericeous, especially when developing fur-like, thin; columella 5.5–7 mm long, fibrous after dehiscence. Seeds (6–)8.5–9(–12) mm long, (4–)7–8(–9.5) mm wide, (4–)5–7(–8) mm high, widened seeds 8–9 mm long, 8–9 mm wide, 6.5–7 mm high.

    Distribution — S Southwestern and Peninsular Thailand, N Malay Peninsula.

    Habitat & Ecology — Evergreen forest; soil: limestone or rarely granite. Altitude: sea-level up to 200(–457) m. Flowering and fruiting: more or less throughout the year.

    Notes — 1. Typical for the species are the enlarged sepals of the calyx when fruiting, the sericeous fruit wall and the long straight fruiting pedicels. When not fruiting this species is easily confused with A. subsessilis (see note under latter for differences).

2. Actephila collinsae, also a species with enlarged sepals in fruit, is endemic in SW Thailand and partly overlaps in distribution with A. ovalis in the province Prachuap Khiri Khan (SW Thailand). Actephila ovalis differs from A. collinsae in the leaf apex (obtuse to bluntly acute in A. collinsae versus acuminate to cuspidate in A. ovalis), leaf margin (flat in A. collinsae, wavy in A. ovalis) and fruiting pedicel and sepals (glabrous in A. collinsae versus slightly hairy on the apical pedicel part and basal sepal part outside in A. ovalis).

3. Gagnepain described several Indochinese species (1925: A. macrantha; 1924: A. nitidula, A. pierrei and A. platysepala) with enlarged sepals in fruit. These have not been assessed, but as they were published later than the epithets collinsae and ovalis, a possible synonymisation will not invoke a name change.

9. Actephila stipularis Heijkoop & Welzen

   Actephila stipularis Heijkoop & Welzen, Blumea 62 (2017) 21, fig. 7, map 6. — Type: Coode 5916 (holo L, 2 sheets; iso K), Indonesia, Sulawesi, Luwuk area, Sulawesi Tengah, Biak to Poh road; 0º49’S, 122º47’E.

Shrubs, up to 1.2 m high, monocaul to sparsely branched with 1 or 2 short branches; flowering branches 5–8 mm diam, with brown lenticels. Indumentum: subglabrous on petioles, leaves and sepals. Stipules large, triangular or ovate, c. 8 by 3 mm, glabrous. Leaves: petioles 4–18 mm long, sometimes somewhat sericeous, completely pulvinate, somewhat flattened, becoming corky with age; blade elliptic to sometimes slightly obovate, 13.3–37.5 by 4.9–9.3, 2.7–4 times as long as wide, papyraceous to pergamentaceous, base acute to somewhat obtuse, margin flat, apex gradually tapering into a widely cuspidate apex, tip acute, abaxially sometimes somewhat sericeous on midrib, often glabrous; nerves c.14 per side. Flowers solitary to paired to fascicled on brachyblasts after several seasons, white. Staminate flowers not seen. Pistillate flowers not seen, in fruit: white; sepals ovate, widest just below the middle, c. 5 by 3 mm, apex obtuse; petals c. 1.5 by 1.3 mm, apex slightly emarginate, erose; disc a narrow, thin ring, slightly covering base of sepals in fruit. Fruits c. 2 cm diam, green; pedicels thick, c. 12 by 1.5 mm, slightly wider at apex, straight, glabrous, striate when dry; sepals not enlarging, reflexed; fruit wall smooth, thin, with 6 indistinct lines of dehiscence; columella c. 10 mm long, basally completely covering the disc, slightly covering the base of the sepals, fibrous after dehiscence. Seeds c. 10 mm long, c. 8 mm wide, c. 7 mm high; widened seeds not seen.

    Distribution — Sulawesi (known from type only).

  A. stipularis: triangles (Sulawesi); A. trigogyna: dots (New Guinea).

    Habitat & Ecology — Forest in steep valleys. Soil: upraised coralline limestone. Altitude: 300 m. Flowering and fruiting: October.

    Note — Typical for the species are the large stipules of c. 8 by 3 mm, in combination with large leaf blades and big fruits on sturdy, thick, short pedicels. The petioles are wide and appear to be flattened. The fruits are completely smooth on the surface and lines of dehiscence are indistinct.

10. Actephila subsessilis Gagnep.

    Actephila subsessilis Gagnep., Bull. Soc. Bot. France 71 (1924) 569; in Lecomte, Fl. Indo-Chine 5 (1927) 534; Heijkoop & Welzen, Blumea 62 (2017) 21, map 7. ― Syntypes: Bon 667 (P, 2 sheets), [Vietnam,] Tonkin, région de Ninh-binh, Hao-nho; Bon 2102 (P), [Vietnam,] Tonkin, région de Hanoï, Kiên-khé, dans les monts Dong-hâm; Bon 5726 (P, 3 sheets), [Vietnam,] Prov. Thanh-hoa, Mâk-són; Eberhardt 4263 (P, 2 sheets), [Vietnam,] Prov. Hoa-binh, Mai-chau; Pételot 1172 (P), [Vietnam,] Cho-ganh; Pételot 1173 (P, 2 sheets), [Vietnam,] Cho-ganh; Pételot 1275 (P, 3 sheets; one noted as 127 in JSTOR website), [Vietnam,] Cho-ganh; Poilane 1658 (P, 2 sheets), [Vietnam,] Annam, Thanh-hóa (collector numbers of Pételot not mentioned by Gagnepain, 1924).

    Actephila excelsa (Dalzell) Müll.Arg. var. acuminata Airy Shaw, Kew Bull. 26 (1972) 209; Whitmore, Tree Fl. Malaya 2 (1973) 52. — Type: Ridley s.n. (holo K), Malaysia, Selangor, Batu caves.

Shrubs to trees; flowering branches with brown lenticels, puberulous, 1.2–3 mm diam. Indumentum: most parts subglabrous. Stipules triangular or ovate, 2–3 by 0.8–1 mm, outside somewhat sericeous when young. Leaves: petioles 1–6(–8) mm long, completely pulvinate, puberulous when young, glabrescent; blade elliptic, 5.6–15 by 2.1–5.2 cm, 2.3–3.7 times as long as wide, papyraceous, base obtuse or acute, margin flat, apex cuspidate to gradually tapering into a widely cuspidate apex, abaxially basally somewhat puberulous on blade and venation; nerves 9–15 per side. Flowers solitary or paired, probably forming small brachyblasts when older. Staminate flowers unknown. Pistillate flowers c. 4 mm diam, yellow; pedicels 12–30 mm long, widening towards apex, slightly sericeous towards apex; sepals ovate to elliptic or diamond shaped, widest just below the middle, c. 2–3 by 1–1.5 mm, apex obtuse; petals 0.8–1 by 0.4–0.8 mm, apex truncate to rounded, erose; disc a thin, narrow ring, slightly covering the base of the sepals in fruit; ovary c. 0.8 by 1.1 mm, glabrous, stigmas c. 0.5 mm long. Fruits green, c. 15 mm diam, pendent; pedicels 17–32 by 0.5–1 mm, somewhat curved, glabrous to sericeous; sepals not elongating, reflexed; fruit wall slightly knobbly outside and with a slightly elevated reticulate venation pattern, glabrous (to slightly sericeous), thin; columella short, sturdy, 4–6 mm long, basally completely covering disc and base of sepals, fibrous after dehiscence. Seeds 7–9.5 mm long, 6–7.5 mm wide, 5–6 mm high, widened seeds c. 8 mm long, 9.5–10.5 mm wide, 6–7 mm high.

    Distribution — Thailand, Vietnam, Malay Peninsula.

    Habitat & Ecology — Only found on limestone. Altitude: 50–550 m. Flowering: April, July; fruiting: September, December, January.

    Note — This species is vegetatively easily confused with A. ovalis, also present in the Malay Peninsula. Differences are the wavy instead of straight leaf blade margin of A. ovalis, and the accrescent calyx in A. ovalis up to 10–22 by 9–13 mm (against c. 2 by 1.4 mm in A. subsessilis).

11. Actephila trichogyna Airy Shaw

    Actephila trichogyna Airy Shaw, Kew Bull. 25 (1971) 499; Kew Bull., Addit. Ser. 8 (1980a) 23; Heijkoop & Welzen, Blumea 62 (2017) 23, map 6. — Type: Brass 8417 (holo L, 2 sheets; iso A, BRI), Papua New Guinea, Western Division, Tarara, Wassi Kussa River.

Trees, 4-5 m high; flowering branches 1-1.75 mm diam, lenticels indistinct, apically puberulous. Indumentum: most parts subglabrous, simple hairs. Stipules triangular or ovate, 1.6–2 by 0.8–1 mm, glabrous. Leaves alternate to subopposite at branch ends; petioles 2–4 mm long, sericeous when young, completely pulvinate; blade elliptic, 5.4–9 by 2.9–4.1 cm, 1.9–2.2 times as long as wide, papyraceous, base acute or obtuse, margin flat, apex obtuse to somewhat acuminate, abaxially somewhat sericeous on midrib when young; nerves 10–13 per side. Flowers green, solitary or paired; sepals: margin membranous, apex obtuse; disc a wide, somewhat fleshy ring. Staminate flowers c. 3 mm diam; pedicel c. 5 mm long, not widening towards apex, glabrous; sepals elliptic, widest in the middle, 1.1–1.2 by c. 1 mm, outside slightly sericious, inside glabrous; petals not seen; disc substantially covering base of sepals; stamens: filaments c. 2 mm long; pistillode not seen, covered by a bundle of hairs. Pistillate flowers not seen; observations from developing fruit: sepals elliptic to somewhat obovate, widest just above to in the middle, 3.5–4 by c. 2 mm, sericeous on both sides; petals 1–1.2 by c. 0.4 mm, apex rounded, erose; disc substantially covering base of sepals in fruit; ovary strongly sericeous. (Developing) Fruits immature globose, mature probably subglobose and flattened; pedicels 13–20 by 0.5–1 mm, slightly widening towards apex, often slightly sericeous, sometimes glabrous, slightly striate when dry; sepals not elongating; wall strongly sericeous, thin; columella not seen. Seeds not seen.

    Distribution — Papua New Guinea (Western Division).

  A. trigogyna: dots (New Guinea); A. stipularis: triangles (Sulawesi).

    Habitat & Ecology — Melaleuca leucadendron swamp-forest. Flowering (and starting to fruit): December.

    Note — Typical for the species are the small leaf blades up to 9 by 4.1 cm, and the short delicate petioles (up to 4 mm long). The fruits are small, but this is probably due to immaturity. Developing fruits resemble A. ovalis in being heavily sericeous with a wide fleshy disc underneath, covering the base of the sepals. However, the sepals are not enlarged and the general leaf blade and stipule shape does not resemble A. ovalis. Staminate flowers have a bundle of hairs instead of a pistillode, it remains unclear whether a pistillode is present underneath.

Actephila spec.nov.? (BW (Koster) 4344)

Heijkoop & Welzen, Blumea 62 (2017) 23.

Shrub, c. 1.75 m high; flowering branches 3–5 mm diam. Indumentum absent. Stipules triangular or narrowly ovate, c. 4 by 2 mm, glabrous. Leaves petioles 13–23 mm long, pulvinate at base, with corky edges when older; blade slightly obovate, 21–29 by 6.4–7.6 cm, 3.3–3.8 times as long as wide, pergamentaceous, base attenuate, tapering into petiole, margin flat, apex acute to somewhat acuminate; nerves 15–18 per side, becoming inconspicuous near margin. Flowers on axillary brachyblasts. Staminate flowers (in bud) white with purple tinge; sepals ovate, apex obtuse, margin membranous, glabrous on both sides; petals 5; disc a complete ring, somewhat lobed, surrounding stamens, folded upwards at the edges; stamens with filaments connate at base. Pistillate flowers, fruits and seeds unknown.

    Distribution — New Guinea (Indonesian Papua: Sowi-Rendani, near Manokwari).

    Habitat & Ecology — Locally rather common in primary forest on clayey soil. Altitude: c. 150 m. Flowering: July.

    Note — The specimen strongly resembles A. lindleyi. However, the stipules are larger, the blade is slightly obovate with an elongated attenuate base and a short, acute apex, a shape not observed in A. lindleyi specimens. Moreover, this specimen has more secondary veins than the A. lindleyi specimens studied. More specimens are needed to certify the status of this specimen as only young staminate flowers were seen.

Excluded species

    Actephila aurantiaca Ridl., Bull. Misc. Inform. Kew (1923) 360; Fl. Malay Penins. 3 (1924) 197. — Type: Ridley s.n., Feb. 1917 (holo K), [Malaysia,] Kelantan, Kelantan River, Chaning. = Trigonostemon aurantiacus (Kurz ex Teijsm. & Binn.) Boerl.

    Actephila flaviflora Lauterb. & K.Schum. in K.Schum. & Lauterb., Fl. Schutzgeb. Südsee (1900) 388. — Syntypes: Hollrung 481 (K), Kaiser Wilhelmsland (= Papua New Guinea), Constantinhafen, Gagar Fluss (Gagar River); Lauterbach 957 (not seen), Kaiser Wilhelmsland, Mittellauf Gogol Fluss (central part Gogol River); Lauterbach 2882 (as 882 in K), Kaiser Wilhelmsland, Nurufluss (Nuru River). = Phyllanthus flaviflorus (Lauterb. & K.Schum.) Airy Shaw (Kew Bull., Addit. Ser. 8, 1980a: 22).

    Actephila megistophylla Quisumb. & Merr., Philipp. J. Sci. 37 (1928) 158. — Lectotype (designated by Kulju & Welzen 2005): BS (Ramos & Edaño) 45466 (holo A; iso K, NY, US), Philippines, Luzon, Tayabas Prov., Casiguran = Cleidion megistophyllum (Quisumb. & Merr.) Airy Shaw (Kew Bull. 25, 1971: 528; Kulju & Welzen, Blumea 50, 2005: 215).