Actephila excelsa (Dalzell) Müll.Arg.

Heijkoop, M. & Welzen, P. C. van, 2017, A revision of the genus Actephila (Phyllanthaceae) in the Malesian region, Blumea 62 (1), pp. 7-25 : 14-17

5. Actephila excelsa (Dalzell) Müll.Arg. — Map 3

Actephila excelsa (Dalzell) Müll.Arg. (1863) 78; (1866) 222; Hook.f. (1887) 282; Pax & K.Hoffm. (1922) 191; Ridl. (1924) 196; Gagnep. (1927) 535; M.R.Hend. (1939) 68; Backer & Bakh.f. (1963) 470; Airy Shaw (1972) 209; Whitmore (1973) 52; Airy Shaw (1975) 25;Chayam. (2005) 35, f. 5. — Anomospermum excelsum Dalzell (1851) 228. — Actephila excelsa (Dalzell) Müll.Arg.var. genuinum Pax & K.Hoffm.(1922) 192,nom.inval. — Actephila excelsa (Dalzell) Müll.Arg. var. excelsa : Airy Shaw (1972) 209. — Type: Dalzell s.n. (holo K barcode K000186757), India, Syhadrensi?

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

Savia actephila Hassk. (1844) 243; Baill. (1858) 571. — Actephila javanica Miq.(1859) 356; Müll.Arg. (1866) 222;J.J.Sm. (1910) 46;Ridl. (1924) 196; Backer & Bakh.f. (1963) 470; Airy Shaw (1980a) 22; (1982) 4; Chayam. (2005) 38, f. 6. — Actephila excelsa (Dalzell) Müll.Arg. var. javanica (Miq.) Pax & K.Hoffm.(1922) 192; Merr.(1923) 390;(1929) 139; Airy Shaw (1972) 209; Whitmore (1973) 52; Airy Shaw (1975) 25; (1980a) 22; (1981) 248; (1983) 3. — Lectotype (designated here): Blume 1670 (L), Java.

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

[ Savia zeylanica Baill. (1858) 571, nom. nud. —] Actephila zeylanica Baill. ex Müll.Arg.(1863) 77; (1866) 221. — Actephila excelsa (Dalzell) Müll.Arg. var. zeylanica (Baill. ex Müll.Arg.) Pax & K.Hoffm. (1922) 192. — Type: Thwaites 2532 (G-DC, P 4 sheets), Ceylon (= Sri Lanka).

Actephila bantamensis Miq.(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.(1863) 77;(1866) 221. — Type: Zollinger 512 (holo B lost; iso perhaps fragment in G-DC as Zollinger s.n.), [ Indonesia,] Java .

Actephila thomsonii Müll.Arg. (1865) 65; (1866) 222. — Actephila excelsa (Dalzell) Müll.Arg.var. thomsonii (Müll.Arg.) Pax & K.Hoffm. (1922) 192. — Type: G. Thomson s.n. (not located), India, Maisor & Carnatic.

Fig. 4 Actephila emarginata Heijkoop & Welzen ( Euphorbiaceae ). Habit with typical emarginate bases Brass 29350, L). ― Drawing by Jan van Os, 2016

Actephila gigantifolia Koord. (1897) 579, 625 (description); Pax & K.Hoffm. (1922) 194. — Type: Koorders 16937 (holo L), [ Indonesia,] Celebes, bivak Totok .

Actephila minahassae Koord. (1897) 579, 625 (description); Pax & K.Hoffm. (1922) 195. — Type: Koorders s.n. (not found), [ Indonesia,] Celebes, Ratahan .

Pimeleodendron dispersum Elmer (1908) 308 ( ‘ dispersa ’); Pax & K.Hoffm. (1912) 56 (excluded species). — Actephila dispersa (Elmer) Merr. (1909) 276. — Type: Elmer 7245 (holo PNH lost; iso K 2 sheets, L), Philippines, Leyte, Palo.

Actephila gitingensis Elmer (1910) 903; Airy Shaw (1975) 25; (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 (1910) 903; Merr. (1923) 390; Pax & K.Hoffm. (1922) 193; Airy Shaw (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 (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. (2009) 717, f. 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, dbh 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 Peninsula, Sumatra, Java, Borneo, the 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 el- liptic 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 speci-

mens 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 spe- cies distribution: Sumatra, Java, Borneo and the Philippines. PPI 6964 ( Philippines) is also special because of the some- what 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 , 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 acute and the ovary is densely hairy instead of glabrous. Therefore, A. puberula (with A. excelsa var. puberula (Kurz) Pax & K.Hoffm. as syno- nym) 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 than sometimes slightly smaller leaf blades (with consequently slightly fewer secondary nerves).

Figures

Map 3 Distribution of Actephila excelsa (Dalzell) Müll.Arg.