|
Malesian Euphorbiaceae Descriptions |
|
Dressler, S. 1999. Revision of the genus Cleistanthus (Euphorbiaceae) in the Philippines. Blumea 44: 109148.
Goto on this page:
Cleistanthus Hook.f. ex Planch. in Hook., Icon. PI. 8 (1848) ad t. 779. Kaluhaburunghos L. ex Kuntze, Revis. Gen. PI. (1891) 607. Type: Cleistanthus polystachyus Hook.f.
Nanopetalum Hassk., Verslagen Meded. Afd. Natuurk. Kon. Akad. Wetensch. 4 (1855) 140. Type: Nanopetalum myrianthum Hassk. [= Cleistanthus myrianthus (Hassk.) Kurz].
Lebidiera Baill., Etude Euphorb., Atlas (1858) 50, t. 27, f. 1-4. = Amanoa sect. Lebidiera Baill., Etude Euphorb. (1858) 581. Type: Lebidiera ferruginea Baill. [= Cleistanthus ferrugineus (Thwaites) Mόll.Arg.].
Stenoniella Kuntze in T. Post & Kuntze, Lex. Gen. Phan. (1903) 535; nom. nov. for Stenonia Baill., Etude Euphorb. (1858)578, (non Endl. 1847). Type: Stenonia boiviniana Baill. [= Cleistanthus stenonia (Baill.) Jabl.].
Leiopyxis Miq., Fl. Ned. Ind., Eerste bijv. (1861) 445. Type: Leiopyxis sumatrana Miq. [= Cleistanthus sumatranus (Miq.) Mόll.Arg.].
Lebidieropsis Mόll.Arg., Linnaea 32 (1863) 79. Type: Lebidieropsis collina (Roxb.) Mόll.Arg. [= Cleistanthus collinus (Roxb.) Benth.].
Schistostigma Lauterb.,Fl. Schutzgeb. Sόdsee, Nachtr. (1905) 299. Type: Schistostigma papuanum Lauterb. [= Cleistanthus papuanus (Lauterb.) Jabl.].
Godefroya Gagnep., Bull. Soc. Bot. France 70 (1923) 435. Type: Godefroya rotundata (Jabl.) Gagnep. [= Cleistanthus rotundatus Jabl.].
Paracleisthus Gagnep., Bull. Soc. Bot. France 70 (1923) 499. Lectotype (Wheeler, 1975: 537): Paracleisthus subgracilis Gagnep. [= Cleistanthus sumatranus (Miq.) Miill.Arg.].
[Zenkerodendron Gilg ex Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 48; in syn., nom. nud.; authentic species: Zenkerodendron bipindense Gilg ex Jabl. (= Cleistanthus bipindensis Pax).]
Trees or shrubs, monoecious,branches scattered lenticellate. Leaves simple, alternate, stipulate, entire, membranous to coriaceous, petiolate; petiole subterete, (slightly) thicker than midrib and wrinkled when dry; brochidodromous venation, secondary veins becoming weaker distally towards theirlooping. Inflorescences axillary, glomerulate, sometimes on leafless or small-leaved branches, glomerules male or female or both. Calyx 5-lobed, mostly persistent, lobes valvate, tube shortly obconic. Petals 5, persistent, small, free. Disc annular, saucer-shaped in staminate flowers and divided in pistillate flowers into an outer saucer-shaped one, lining the receptacle, and an inner cup-shaped to tubularonewith irregular margin, covering the ovary (or at least its base) and tearing into lobes during fruit development. Stamens 5, filaments connate into a column, free above, spreading, terete to flattened, tapering towards the apex, anthers extrorse, bilobed at the base, pistillode small, trifid or trilobed, borne at the apex of the column. Ovary 3-locular, ovules 2 in each cell; stigmas 3, subconnate below, bifid. Capsules 3-locular, subglobose or depressed, sessile or pedicelled, sometimes stipitate within the calyx, dehiscing apically loculicidal and completely septicidal, hence the valves falling apart, leaving the receptacle and columella; dehiscence is mechanical, caused by the expansion of the internal endocarp tissue, additionally septae splitting during this process; endocarp bony, inside fibrous, each locule 2-seeded or frequently 1-seeded by abortion. Seed exarillate.
|
1a. |
Leaves glabrous beneath (exceptionally scattered hairs present) |
|
|
1b. |
Leaves hairy beneath(at least on the main veins) |
|
|
Flowers (and fruits) clearly pedicellate (pedicel > 5 mm long) |
||
|
2b. |
Flowers and fruits (sub)sessile |
|
|
Leaves with conspicuously prominent, fine, dense, reticulate venation, greyish brown beneath. Many flowers in axils of normal leaves |
||
|
3b. |
Leaves with inconspicuous venation, dark coloured (blackish) beneath. Flowers on short axillary shoots (brachyblasts) |
|
|
Capsules big, > 15 mm diam. Stipules 510 mm long, long persistent. Ovary glabrous. Leaves often conspicuously glaucescent beneath |
||
|
4b. |
Capsules smaller, < 15 mm diam. Stipules < 5 mm long. Ovary densely hairy* [*cf. Cleistanthus spec. A: ovary presumed to be glabrous] |
|
|
Secondary veins more than 9 pairs. Capsules stipitate (24 mm) and subsessile to shortly pedicellate (24 mm), densely brownish tomentose. Petiole mostly > 7 mm, scattered pilose to glabrous |
||
|
5b. |
Secondary veins less than 9 pairs. Capsules not stipitate and sessile, scattered pilose to glabrous. Petiole mostly < 7 mm, glabrous |
|
|
Stipules 24 mm long, triangular. Veins conspicuously prominent beneath.Flowers on smaller leaved branchlets |
||
|
6b. |
Stipules up to 1.5 mm long, scale-like or triangular. Veins faintly prominent beneath. Flowers axillary on normal leaved or short, leafless, axillary, spike-like branchlets |
|
|
Leaves yellowish or greenish when dry; base obtuse, slightly attenuate. Secondary veins not conspicuously long running towards apex |
||
|
7b. |
Leaves (olive) brownish when dry; base acute to round, rarely obtuse. Secondary veins with a rather acute angle of divergence and long running towards apex before arching |
|
|
Leaves conspicuously narrow-elliptic or narrow-ovate (nearly linear). Long drooping branches. Flower glomerules usually in the axils of normal leaves |
||
|
8b. |
Leaves not extraordinarily narrow. Branches short, not drooping. Flower glomerules on differentiated, often smaller leaved to leafless, terminal or axillary branchlets (like 'pearls-on-a-string') |
|
|
Capsule > 15 mm diam. Stipules conspicuous, 510 mm long, long persistent. Leaves conspicuously glaucescent beneath (see however C. everettii with 29 mm long stipules but not conspicuously glaucescent leaves) |
||
|
9b. |
Capsule < 15 mm diam.Stipules < 5 mm long, not conspicuous, not long persistent (if so, see C. everettii). Leaves not conspicuously glaucescent beneath |
|
|
Leaves finely and closely appressed puberulous to sericeous beneath (lens!). Ovary and capsule glabrous |
||
|
10b. |
Leaves with erect hairs beneath. Ovary (and later capsule) ± hairy |
|
|
Leaves with coppery indumentum beneath. Secondary veins mostly 58 pairs . |
||
|
11b. |
Leaves with light to golden indumentum beneath. Secondary veins mostly 1016 pairs |
9. Cleistanthus myrianthus |
|
Capsules not stipitate |
||
|
12b. |
Capsules stipitate |
|
|
Branchlets conspicuously long, light-coloured pilose, also main veins. Leaves greenish-brown when dry |
||
|
13b. |
Branchlets fulvo-pubescent to -puberulous. Leaves pale green to olive when dry |
|
|
Leaves linear-oblong. Stipules minute, < 1 mm long, very inconspicuous |
||
|
14b. |
Leaves elliptic. Stipules 1 mm and longer |
|
|
Petioles 1.54 mm long. Capsules < 1 cm diam. Leaves mostly smaller (< 9 cm long) |
||
|
15b. |
Petioles 48 mm long. Capsules > 1 cm diam. Leaves mostly larger (> 8 cm long) |
|
|
Glomerules few-flowered (< 10 flowers) |
||
|
16b. |
Glomerules many-flowered (> 10 flowers) |
|
|
Leaf base attenuate, conspicuously decurrent along petiole. Leaves dark reddish brown when dry, blade elliptic to ovate. Stipules > 1 mm long. Sepals fulvo-strigose to subglabrous |
||
|
17b. |
Leaf base acute, but not decurrentalong petiole. Leaves greenish when dry, blade oblong. Stipules minute, < 1 mm long. Sepals glabrous |
|
|
Leaves glabrous, rarely with some hairs, brown to greenish-grey when dry, often bullate. Secondary veins mostly up to 9 pairs. Capsules > 1 cm diam., densely brownish tomentose, subsessile to shortly pedicellate (24 mm) |
||
|
18b. |
Leaves rusty pubescent to pilose, rarely subglabrous, dark reddish brown when dry. Secondary veins mostly less than 9 pairs. Capsules < 1 cm diam., scattered pilose to glabrous. Inflorescence glomerules conspicuously rufo-villose |
Cleistanthus angustifolius Merr.,Philipp. J.Sci., Bot. 7(1912) 386; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 52; Merr., Enum. Philipp. Flow. PI. 2 (1923) 419; Salvosa, Lex. Philipp. Trees (1963) 92; Airy Shaw, Enum. Euphorb. Philipp. Is. (1983) 14; S.Dressler, Blumea 44 (1999) 113, map 1. Lectotype (designated by Dressler 1999): C.M. Weber 1562 (holoA; iso E, F, G, GH, K, NY, P 3x, US), Philippines, Luzon, Prov. Cagayan, Abulug River.
Scandent shrub or tree? Branches often long, of drooping appearance, glabrous, rarely scattered albo-pilose when young. Stipules very inconspicuous, triangular, c. 1 by 0.51 mm, glabrous, scaly. Leaves: petiole subterete, darker coloured than midrib in dry state (blackish), glabrous, 36 by 0.51 mm; lamina very narrow ovate to narrow ovate-elliptic or narrow lanceolate (almost linear), 35130 by 630 mm, 3.69.6 times as long as wide, chartaceous, base roundish, rarely obtuse, margin entire, apex long acute to acuminate, acumen not discernible, glabrous on both sides, greyish brown to brownish olive above, greyish green to brownish grey beneath when dry, sometimes glaucescent beneath; venation obscure above, very faintly prominent beneath, secondary veins in 58 pairs, long running towards apex before looping, sometimes with tertiary arches, irregular reticulate areolation. Inflorescences: few-flowered glomerules (up to c. 5 flowers) in the axils of normal leaves, rarely on short, spike-like shoots; bracts small, triangular, c. 1.5 by 11.5 mm, glabrous to albopilose, with sericeous margin, and/or hairy midvein. Flowers (not seen) sessile, 45 mm diam. Sepals (narrow-)triangular, c. 2 by 11.5 mm, albo-pilose outside. Infructescences with 1 or 2 fruits per glomerule. Fruits sessile, rarely minutely stipitate (c. 1 mm), obtusely triangular and deeply 3-lobed from above or tricoccous-subglobose, concavely depressed at apex, 56 mm tall, c. 8 mm diam., glabrous, basal and in sutures (grooves) with scattered pilose hairs, dark brown when dry. Seeds semiglobose to heart-shaped, ventrally with median hilum, dorsally slightly keeled, 67 by 4.55.5 by 33.5 mm; testa smooth, slightly lineate with occasional inconspicuous warts, brown.
Distribution Philippines (Luzon).
l
= C. angustifolius; n
= C. erycibifolius; p
= C. glaber; q = C.
isabellinus
Habitat & Ecology On river.
Notes The main characteristics are: conspicuously narrow leaves (nearly linear, narrow ovate to narrow elliptic), long drooping, glabrous branches, glabrous, chartaceous leaves with indistinct, i.e., hardly prominent venation, 58 secondary veins, sessile fruits, which are sparsely pilose and become subglabrous.
Only 4 collections of this species could be traced and all are in fruit.
It might well be that this species merely represents a very narrow leaved (rheophytic?) form of C. sumatranus. Unfortunately, there are no fieldnotes. The type collection was collected at Abulug River, Cagayan Province, Luzon. Two other collections are both annotated with Cagayan Province only. All three were collected in January 1912. It is possible that all three originate from the same plant. An additional collection from Zambales Province, Luzon, has larger leaves which are already somewhat intermediate to C. sumatranus. However, the plants display the drooping habit.
I (= Dressler 1999) provisionally retain the species rank here. The taxon is separated from C. sumatranus by the combination of its distinct leaf shape and drooping habit.
Cleistanthus bridelifolius C.B.Rob., Philipp. J. Sci., Bot. 3 (1908) 191; Philipp. J. Sci., Bot. 6 (1911) 323; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 25; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 419; Salvosa, Lex. Philipp. Trees (1963)92; Airy Shaw, Kew Bull., Add. Ser. 4 (1975) 78; Kew Bull. 33 (1978) 51; J.A.R. Anderson, Check List Trees Sarawak (1980) 181; Airy Shaw, Kew Bull. 36 (1981) 280; Kew Bull. 37 (1982) 13; Enum. Euphorb. Philipp. Is. (1983) 14; Whitmore, Tree Fl. Indon.,Checkl. Sumatra (1986) 80; Tree Fl. Indon., Checkl. Sulawesi (1989) 46; Tree Fl. Indon., Checkl. Kalimantan 1 (1990) 123; Keίler & Sidiyasa, Trees Balikpapan-Samarinda Area, Tropenbos ser. 7 (1994) 126 in clavi; S.Dressler, Blumea 44 (1999) 115, map 2. Lectotype (designated by Dressler 1999): FB (Klemme) 7064 (holo US), Philippines, Luzon, Prov. Cagayan, San Vicente. [Note: Airy Shaw (1978) suspected the type to be in NY. I did not receive a specimen from there.]
Cleistanthus pallidus (Thwaites) Mull. Arg. var. subcordatus J.J.Sm. in Koord. & Valeton, Bijdr. Boomsoort. Java 12 (1910) 304. Cleistanthus subcordatus (J.J.Sm.) Jabl. in Engl., Pflanzenr. IV.147.viii (1915)22; Backer & Bakh.f., Fl. Java 1 (1964) 474 in clavi. Type: Teijsmann HB 1741 (BO?), Java, Madoera, M. Gegu - protologue:"alleen verzameld in de res. Banten, afd. Tjaringinbij Tjemara op 10-200 m zeehoogte". [Note: Although I have not seen the actual type specimen, I am convinced of the conspecificity since material identified by J.J.Smith in L represents C. bridelifolius.]
Shrub to tree, up to 15 m high, reported with 6 m high bole and with up to 35 cm dbh. Bark smooth to rough, brown, inner bark reddish brown, cambium pinkish; sapwood yellow. Branches fulvous-pubescent to -puberulous when young, later becoming glabrous, triangular stipules sometimes conspicuously densely concentrated and long persistent on the terminal branches often after the leaves have fallen. Stipules narrow to broadly ovate-triangular, subulate, 13.5 by 11.5 mm, glabrous to puberulous. Leaves: petiole subterete, densely pubescent to scattered pilose, rarely glabrous, 1.54 by 11.4 mm; lamina elliptic to lanceolate, rarely slightly ovate, 4090(130) by 1030 mm, 2.34.2 times as long as wide, chartaceous, base acute, roundish to obtuse, sometimes slightly cordate, margin entire, apex acute to acuminate, acumen up to 15 mm long, upper surface glabrous, only single hairs at midrib, or near base and margin, (scattered) pilose to glabrous beneath, densely hairy on base and major veins, rarely fully glabrous, both surfaces greyish green to olive when dry; venation: secondary veins in 611(12) pairs, (sub)prominent beneath, obsolete above, areolation indistinct. Inflorescence: few-flowered glomerules with up to c. 8 flowers, axillary on normal-leaved branches; bracts broadly ovate-triangular to triangular, c. 1 by 11.5 mm, densely whitish pubescent. Flowers sessile to shortly pedicellate, fragrant; pedicel 12(3) mm long, pubescent, rarely glabrous, 3.55 mm diam. Sepals triangular, 1.52 by 11.5 mm, pale green whitish pubescent to subglabrous outside. Petals whitish, spathulate to broadly obovate, lobulate at apex, 0.71 by 0.60.7 mm. Disc glabrous, sometimes pilose, in staminate flowers and outer one in pistillate flowers saucer-shaped, 11.5 mm diam., inner one cup-shaped with erose margin, 0.81.2 mm tall, glabrous to whitish pilose outside. Stamens yellowish; staminal column 11.2 mm long; free part of filaments up to c. 1 mm long; anthers ovoid, c. 0.5 by 0.20.3 mm; pistillode conico-ovoidal, whitish pilose, 0.50.7 by 0.20.4 mm. Ovary ovoid to globose, 11.5 by 0.81.2 mm, densely whitish strigose; stigmas 0.51(1.5) mm long, apically bifid, lobulate. Infructescence with 1 or 2 (also 3?) fruits. Fruits sessile, triangular seen from above, shallowly 3-lobed in outline, concavely depressed at central apex, 57 mm tall, 79 mm diam., light to yellowish green turning dark purple with greenish base, yellowish pilose to pubescent, rarely nearly glabrous. Seeds semi-ovoid to heart-shaped, sometimes abaxially keeled, 34 by 3.54.5 by 1.53mm; testa smooth to rugulate, dark brown.
Distribution Peninsular Malaysia? (fide Whitmore, 1986), Sumatra, Borneo, Philippines, Java (fide Whitmore, 1986), Celebes, Flores (fide Airy Shaw, 1982).
l
= C. bridelifolius; n = C. everetii
Habitat & Ecology Open or low statured primary or secondary lowland forests; at altitudes up to 800 m. Reported from ultrabasic, igneous-derived soils, black stony soil, or coral limestone.
Notes The diagnostic characters of C. bridelifolius are: the pubescence of branches, petiole, inflorescence glomerules axillary; the stipules glabrous to scattered puberulous; petiole 24 mm long; inflorescences axillary; capsules sessile, < 1 cm diam. The leaves are smaller than those of C. everettii, to which it is similar; the distinguishing characters are petiole length and capsule size. Further studies are needed in order to show whether there is a real discontinuity between these species.
Ironically, this species does not particularly resemble Bridelia in its foliage. Robinson did not state in the protologue why he had chosen this name. There are other species which more closely resemble the leaves of certain Bridelia species of the sect. Cleistanthoideae. A flowering Cleistanthus vestitus Jabl. e.g. recalls B. insulana Hance or B. glauca Blume. The untrained eye has to examine the number of ovary locules or styles to trace the generic affiliation.
Airy Shaw recognises two varieties. His var. calcicola differs from the typical one in less abruptly and distinctly caudate leaf-apices, a rather rectangular angle of divergence of the secondary veins, pedicellate staminate flowers, glabrous sepals and its occurrence on limestone. This combination of characters is, at least in Philippine material, not consistent. Airy Shaw's (1978: 51) statement concerning an even higher ranking of this entity cannot be supported. Ecological field studies are necessary in order to decide whethera supposedly calcicole variety should be recognised at all.
Cleistanthus decurrens Hook.f., Fl. Brit. India 5 (1887) 278; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 33; Ridl., Fl. Malay Penins. 3 (1924) 191;Gage, J. Asiat. Soc. Bengal 75 (1936) 508; Henderson, J. Malayan Branch Roy. Asiat. Soc. 17 (1939) 69; Airy Shaw, Kew Bull. 26 (1972) 236; Whitmore, Tree Fl. Malaya 2 (1973) 81 in clavi; Airy Shaw, Kew Bull., Add. Ser. 4 (1975) 80; J.A.R. Anderson, Check List Trees Sarawak (1980) 181; Airy Shaw, Enum. Euphorb. Philipp. Is. (1983) 14; Whitmore,Tree Fl. Indon., Checkl. Kalimantan 1 (1990) 123; S.Dressler, Blumea 44 (1999) 117, map 3. Kaluhaburunghos decurrens (Hook.f.) Kuntze, Rev. Gen. PI. 2 (1891) 607, nom. illeg. Lectotype (designated by Dressler 1999: 117): Scortechini 1916 (holo K), Peninsular Malaysia, Perak.
Cleistanthus decipiens C.B.Rob.,Philipp. J.Sci.,Bot. 3 (1908) 195; Elmer, Leafl. Philipp. Bot. 4 (1911) 1284; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 33; Merr., Enum. Philipp. Flow. PI. 2 (1923) 420; Salvosa, Lex. Philipp. Trees (1963) 92; Airy Shaw, Enum. Euphorb. Philipp. Is. (1983) 14. Lectotype (designated by Dressler 1999: 117): FB (Clark) 1001 (holo K; iso NY), Philippines, Island Ticao, Niladlaran.
Cleistanthus ovatus C.B.Rob., Philipp. J. Sci., Bot. 3 (1908) 194; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 34; Merr., Enum. Philipp. Flow. PI. 2 (1923) 421; Salvosa, Lex. Philipp. Trees (1963) 92; Airy Shaw, Enum. Euphorb. Philipp. Is. (1983) 15.Lectotype (designated by Dressler 1999: 117): BS (Fenix) 4051 (holo K; iso US), Philippines, Camiguin Island, Babuyanes.
Cleistanthus mattangensis Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 33l; Airy Shaw, 1975; Merr., Bibl. Enum. Born. Pl. (1921) 334; Masam., Enum.Phan. Bornear. (1942) 392. Lectotype (designated by Dressler 1999: 117): Beccari 2040 (holo K; iso FI-B, photo seen), Borneo, Mattang.
Cleistanthus decipiens auct. non C.B. Rob.: Elmer, Leafl. Philipp. Bot. 4 (1911) 1284; Merr., Enum. Philipp. Flow. PI. 2 (1923) 420, p.p. quoad Elmer 12888 & 12988.
Shrub to small tree, up to 5 m high, with a suberect, crooked stem, dbh c. 3 cm. Outer bark yellowish grey, smooth, thin; sapwood yellowish-white; heartwood rose-red. Branches brownish strigose to scattered pilose when young, soon glabrescent. Stipules triangular, often scale-like, 1.52 by 11.5 mm. Leaves: petiole (sub)terete, darker coloured than midrib in dry state, brownish strigose, (3)47 by 12 mm, swollen when fresh; lamina lanceolate, elliptic to (slightly) ovate, 35215 by 1899 mm, 1.83.8 times as long as wide, chartaceous to subcoriaceous, base attenuate, conspicuously decurrent on petiole, margin entire, slightly revolute, apex acuminateto (bluntly) acute, acumen up to 20 mm long, above scattered albo-pilose, glossy dark green, beneath albo-pilose to subglabrous beneath, sometimes strigose near base and at midrib, lighter beneath than above, both surfaces often conspicuously dark reddish brown when dry; venation: secondary veins in (5)711 pairs, prominent beneath, (sub)prominent above, sometimes tertiary arches present, perfect reticulate areolation. Inflorescences: few-flowered glomerules (mostly 3 or 4, up to c. 7 flowers), axillary on normal-leaved branches, very rarely axillary, spike-like, leafless branches bearing the glomerules seen; bracts fulvous, elliptic, ovate or broadly triangular, 1.52.5 by 1.52 mm, membranous, often conspicuously brownish strigose to pilose outside, sometimes glabrous with hairy midvein, with erose margin. Flowers sessile, 34.5 mm diam., pale green. Sepals triangular, 1.52 by 11.5 mm, brownish strigose to subglabrous with some hairs on apex outside. Petals whitish, spathulate, margin irregularly erose to lobulate, c. 1 by 0.50.8 mm. Disc glabrous, in staminate flowers and the outer one in pistillate flowers saucer-shaped, 11.5 mm diam., inner one cup-shaped, 0.50.8 mm tall. Staminal column 11.5 mm long, free part of filaments up to 1 mm long; anthers ovoid, c. 0.5 by 0.20.4 mm; pistillode cylindrical, with widened blunt apex, glabrous, up to 0.7 by 0.3 mm. Ovary (semi)- globose, 11.5 by c. 1.5 mm, densely brownish strigose to pilose; stigmas c. 1 mm long, apical quarter bifidly divided, widened tip, latter lobulate. Infructescences with up to 2 fruits. Fruits sessile, but 46 mm stipitate, obtusely triangular from above, shallowly tri-lobed, more or less rectangular to semiorbicular with concave depression in outline, 711 mm tall, 710 mm diam., reddish green turning black, stipe yellow-green, sparsely pilose to glabrous with some scattered hairs in grooves and at base, shiny reddish brown when dry with prominent reticulum of veins, or tuberculate. Seeds heart-shaped in outline (?, collapsed), 56 by 33.5 by 23 mm; testa smooth to rugulate, light brown.
Distribution Peninsular Thailand, Peninsular Malaysia, Borneo, Philippines, Java (?), Flores (?).
l
= C. decurrens; n = C. vestitus
Habitat & Ecology Primary or secondary forests, often reported from dense shade; at low altitudes to 500 m; very fertile, rocky soil, occasional on limestone (Whitmore 1973).
Notes This species is characterised by the decurrence of the basal leaf margins along the petiole, as well as by its reddish brown leaves in the dried state, its few-flowered glomerules, and its stipitate, nearly glabrous capsules.
Cleistanthus decipiens, described as endemic for the Philippines, is according to Jablonsky (1915)'s key distinguished from C. decurrens by the shorter stipitate capsules (< 4 versus 610 mm) and more or less acuminate (versus rotundate or obtusely acuminate) leaves. However, material from Peninsular Malaysia and Borneo shows all intergradations in leaf tips. Airy Shaw annotated the isotype in K as "very near C. decurrens but did not place the name into the synonymy of the latter. This is done here now. In the protologue of C. decipiens Robinson (1908) already stated that this species is "very similar in gross characters to C. ovatus, but quite distinct, especially in its venation". It is said to differ from the latter by having elliptic or oblong leaves (versus ovate ones) and by curving secondary veins (versus straight ones) which do not submarginally divide forming distinct tertiary loops as it was deemed to be typical in C. ovatus. Having examined some more material I have to state that this distinction is not consistent, but merely due to different developmental stages and shapes of the leaves. In fact, the tertiary loops are often present in all leaves, but sometimes not very clearly developed or very small and close to the margin.
The reference to Flores is based on Verheijen 3077, Schmutz 4361 (both L), which are identified as cf. decurrens. There is also a collection of C. decurrens in Leiden (Boerlage s.n., L no. 909.77-179, 909.76-164) which is filed among Javanese material, but the provenance is a bit doubtful as the label information is hardly legible (I would read: "Eiland Leiden, Boerlage"). Van Steenis-Kruseman (1950: 68) gives Pulau Leiden in the Bay of Batavia as a collecting locality of Boerlage in Sept. 1896.
Cleistanthus erycibifolius Airy Shaw, Kew Bull. 20 (1967) 389; Kew Bull. 27 (1972) 76; Kew Bull., Add. Ser. 4 (1975) 80; Kew Bull. 32 (1978) 383; Whitmore, Tree Fl. Indon., Checkl. Kalimantan 1 (1990) 124; S.Dressler, Blumea 44 (1999) 120, fig. 1, map 1. Type: Schut K.24 (holo K; iso A n.v., BO, L, P n.v., PNH n.v., SING), E Indonesian Borneo, Kutei, River Tiram.
Tree, up to 30 m high, with a clear bole up to 10 m high, up to 25 cm dbh, and a round crown. Outer bark light brown, soft, peeling off; inner bark brown, bark bitter; sapwood yellowish to pale white. Branches glabrous to scattered puberulous, horizontal. Stipules triangular, small, c. 1 by 0.61 mm, glabrous, scale-like, often inconspicuous. Leaves: petiole subterete, dark coloured in dry state (blackish), glabrous; blade elliptic, sometimes slightly obovate, 40130 by 1338 mm, 2.43.9 times as long as wide, subcoriaceous, base acute to obtuse, margin entire, slightly revolute, apex acuminate, acumen 1520 mm long, above glabrous, light brownish grey, conspicuously light brown beneath when dry, beneath whitish or glaucou, glabrous to rarely scattered puberulous on petiole and midrib; venation: inconspicuous on both sides, obscure above, obscure to indistinctly raised beneath, secondary veins in (5)610(11) pairs, secondary arches only slightly weaker than secondary veins, areolation indistinct. Inflorescences: flowers on short, condensed, axillary brachyblasts, these on short axillary shoots or in axils of normal leaves; bracts small, broadly ovate to ovate-triangular, 0.81.5 by 0.51 mm, glabrous to puberulous outside, margin erose, sometimes slightly ciliate, abaxially keeled (midvein). Flowers not seen!; distinctly pedicelled (?), 58 mm diam. (in fruit). Sepals triangular, 23 by 1.51.8 mm, glabrous. Inner pistillate disc cup-shaped, c. 1.21.5 mm tall (in fruit), margin irregular. Stigmas 11.5 mm long, deeply bifid (up to half of their length), tips widened, lobulate. Infructescences seen with up to 2 fruits per brachyblast. Fruits distinctly pedicelled, pedicel 37 mm long, 12 mm diam.; subglobose, depressed, trilobed from above,79 mm tall, 810 mm diam.,glabrous, only sporadic single hairs, especially basal. Seeds bean-shaped, ventrally concavely depressed with medial hilum, 67.5 by 45 by 45 mm; testa smooth, brown.
Distribution Peninsular Malaysia, Borneo, Philippines (Negros).
n
= C. erycibifolius; l
= C. angustifolius; p
= C. glaber; q = C.
isabellinus
Habitat & Ecology Primary forests, locally common; up to 1400 m. Reported from sandy soil, sandstone, or sandy loam.
Notes This only rather recently described but quite characteristic species is here reported for the first time for thePhilippines. Its characteristics are: stoutly pedicelled fruits, which are borne on shortly condensed shoots (brachyblasts), subcoriaceous leaves with an indistinct venation and a conspicuous ochre-brown colourbelow, as well as its general glabrousness in young parts.
Probably most closely related to C. pedicellatus this species is easily distinguishable from that by its thicker leaves, which have an indistinct venation when dried (rarely faintly prominent) and the shorter, but stouter pedicel of the fruits. Non-fruiting collections sometimes recall C. bridelifolius, but seem strange because of their subcoriaceous, glabrous leaves, and their inconspicuous scale-like stipules.
Cleistanthus everettii C.B.Rob., Philipp. J. Sci., Bot. 3 (1908) 194; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 53; Merr., Enum. Philipp. Flow. PI. 2 (1923) 420; Salvosa, Lex. Philipp. Trees (1963) 92; Airy Shaw, Enum. Euphorb. Philipp. Is. (1983) 14; S.Dressler, Blumea 44 (1999) 121, fig. 2, map 2. Lectotype (designated by Dressler 1999): FB (Everett) 7274 (holo K; iso US), Philippines, Negros, Himugaan River.
Cleistanthus samarensis Merr., Philipp. J. Bot. 9 (1914) 475; Enum. Philipp. Flow, PI. 2 (1923) 422; Salvosa, Lex. Philipp. Trees (1963) 93; Airy Shaw, Enum. Euphorb. Philipp. Is. (1983) 16. Lectotype (designated by Dressler 1999); BS (Ramos) 1685 (holo G; iso BO, G, GH, L, NY, P, SING, US), Philippines, Samar, Mt Cauayan.
Cleistanthus acuminatissimus Merr., Univ. Calif. Publ. Bot. 25 (1929) 155; Masam., Enum. Phan. Bornear. (1942) 391; Meijer, Bot. News Bull. Forest Dept., Sabah 7 (1967) 41 in clavi; Bot. Bull. Herb. Forest Dept., Sabah 10 (1968) 232; Airy Shaw, Kew Bull., Add. Ser. 4 122 (1975) 77; Whitmore, Tree Fl. Indon., Checkl. Kalimantan 1 (1990) 122. Lectotype (designated by Dressler 1999): Elmer 20969 (holo PNH, lost, replaced by L; iso BM n.v., BO, BR, DS, GH, K, L, M, NY, P, PH, SING, U, UC), British N Borneo, nearr Tawao (Elphinstone Prov.)
Shrub to small tree, up to 6 m high, up to 20 cm dbh. Heartwood light olivaceous. Branches fulvous-pubescent to -pilose when young, later becoming glabrous, triangular stipules sometimes conspicuously densely concentrated and long persistent on the terminal branches. Stipules ovate- to narrow-triangular, subulate, glabrous to scattered pubescent, 29 by 12 mm. Leaves: petiole subterete, greenish to brownish when dry, pubescent to scattered pilose, 48 by 1.52 mm; lamina lanceolate to elliptic, rarely slightly obovate, 60240 by 2095 mm, 1.94.5 times as long as wide, chartaceous, base acute to obtuse, rarely slightly emarginate, margin entire, apex attenuate to acuminate, acumen 1525 mm long, above deep green, glabrous excepr only single hairs at midrib or near base and margin, beneath light green, (scattered) pilose to glabrous beneath, densely hairy on base and major veins, surfaces drying greyish green, sometimes conspicuously rugose; venation:secondary veins in 912(14) pairs, prominent beneath,obsolete above, areolation indistinct. Inflorescences: few-flowered glomerules with up to 10(15) flowers, axillary on normal-leaved branches, often with conspicuous whitish, dense pubescence; bracts broadly triangular, 2.53 by c. 2.5 mm, whitish pubescent. Flowers sessile c. 56 mm diam. Sepals triangular, (1.5)23 by 11.6 mm, densely, whitish- to yellowish-strigose to -pubescent outside. Petals whitish to light yellow, spathulate to broadly obovate, lobulate at apex, 0.81 by 0.81 mm. Disc in staminate flowers and outer one in pistillate flowers saucer-shaped, 23 mm diam., inner one cup-shaped with erose margin, 1.5(2) mm tall, whitish strigose outside. Staminal column c. 1.5 mm long, free part of filaments up to c. 1 mm long; anthers ovoid, c. 0.8 by 0.5 mm; pistillode conical, whitish strigose to pilose, 0.81 by 0.50.7 mm. Ovary globose, c. 1.5 by 1.5 mm, densely whitish strigose; stigmas 0.71 mm long, apically indistinctly bifid. Infructescences with 1 or 2 (also 3?) fruits. Fruits (sub)sessile (base sometimes c. 1 mm stalked), roundly obtuse-triangular from above, 3-lobed in outline, concavely depressed at apex, 810 mm tall, 1013 mm diam., densely yellowish pubescent. Seeds heart-shaped, sometimes abaxially keeled, 55.5 by 4.55 by 3.54 mm; testa smooth to rugulate, brown.
Distribution Borneo, Philippines.
n = C. everetii;
l
= C. bridelifolius
Habitat & Ecology Primary forests; up to 700 m altitude. Reported from red clay soil.
Notes This species is characterised by pubescent branches, petioles, and inflorescence glomerules, but glabrous (to scattered pubescent) stipules. The axillary inflorescences are oftenconspicuously whitish pubescent. The sessile capsules have a diameter > 1 cm and the leaves are larger than in C. bridelifolius.
Robinson described C. everettii in the same article as C. bridelifolius, and from the protologues it seems to be obvious that it is mainly based on the larger leaves, a longer petiole (58 mm), more secondary veins (1113) and a shortly pedicelled capsule (this could not be confirmed in the type material in K andUS). No qualitative characters could be found which distinguish C. everettii from C. bridelifolius. The distinguishing characters between C. everettii and C. bridelifolius are: petiole length, leaf size, number of secondary veins, and size of the capsules. It seems that C. everettii merely represents a larger version of C. bridelifolius. The distinction between both is occasionally somewhat problematic and I am aware that the gap may appear rather artificial. However, I retain both here as species until further information is at hand: lumping both would render the species enormously variable in leaf size, and one would have to lump other taxa as well for the sake of inherent logic and consistency.
As so often field studies could contribute towards a clarificationofthis taxonomically unsatisfying situation.
The type of C. samarensis (Ramos 1685) was collected from a narrow-leaved specimen of C. everettii. From the protologues of these species it is evident that they differ only by lanceolate (samarensis) vs. elliptic or oblong leaves (everettii). As there are plenty of intermediate collections available, I consider them to be conspecific.
Due to the absence of distinguishing characters C. acuminatissimus is here subsumed under C. everettii which extends the latter's range to Borneo.
Cleistanthus glaber Airy Shaw, Kew Bull. 21 (1968) 366; Meijer, Bot. News Bull. Forest Dept., Sabah 7 (1967) 40 in clavi; Bot. Bull. Herb. Forest Dept., Sabah 10 (1968) 230; Whitmore, Tree Fl. Malaya 2 (1973) 80 in clavi; Airy Shaw, Kew Bull., Add. Ser. 4 (1975) 80; J.A.R. Anderson, Check List Trees Sarawak (1980) 181; Airy Shaw, Enum. Euphorb.Philipp. Is. (1983) 14; Whitmore,Tree Fl. Indon.,Checkl. Kalimantan 1 (1990) 124; S.Dressler, Blumea 44 (1999) 124, map 1. Type: J. Clemens & M.S. Clemens 22236 (holo K; isoA, BO 2x, K, L, P), Borneo, Sarawak, Lundu, Mt Gadin.
Tree, up to 15 m high, with a clear bole up to 6 m high, girth up to 1 m. Outer bark green, flaky to scaly; inner bark white or reddish; cambium white or brown; sapwood pale yellowish-white to brown. Branches glabrous. Stipules scaly, often inconspicuous, triangular, minute, 0.71.2 by 0.51 mm. Leaves: petiole subterete, coloured differently from the blade in dry state (yellowish or blackish), glabrous, 37 (8) by 12 mm; lamina ovate to elliptic, very rarely obovate, 44165 by 1887 mm, 1.83.5 times as long as wide, chartaceous, base obtuse, slightly attenuate, rarely acute, margin entire, apex acuminateto acute, acumen (10)1525 mm long, surfaces glabrous, greenish to yellowish when dry, hardly glaucescent beneath, butreported to be glaucous beneath when living; venation: secondary veins in 49 pairs, prominent beneath, secondary veins rather irregular, perfect reticulate areolation. Inflorescences: few-flowered glomerules (up to c. 10 flowers) axillary (mostly on terminal branches) or on short axillary spikes (up to 4 cm long) with the spike axis sometimes scattered pilose; bracts small, ovate-triangular, c. 1 by 1 mm, pilose outside, midvein glabrous, with erose margin. Flowers sessile, staminate ones slender, c. 4 mm diam., pistillate ones 5(6) mm diam. Sepals triangular, 1.82.5 by 1.41.8(2) mm, brownish (staminate) or greenish (pistillate) when dry, densely to scattered, appressed albo-pilose outside, indumentum becoming scattered in fruit. Petals whitish, obovate to rhomboidal, margin irregularly erose, 0.81 by 0.50.6 mm. Disc in staminateflowers 1.61.7mm diam., outer one in pistillate flowers saucer-shaped, c. 2 mm diam., inner one cup-shaped. Staminal column c. 0.6 mm long (immature); free part of filaments up to 0.7 mm long (immature); anthers ellipsoid, two thecae well separated, 0.50.6 by c. 0.4 mm; pistillode broadly ovoid, hairy, up to 0.5 by c. 0.3 mm. Ovary globose to slightly ovoid, apically terminating into stigmas, 11.2 by 1.21.6 mm, densely hairy; stigmas 1.52 mm long, united at base, apical branches of bifidly divided (upper thirds or quarters), tips papillose, widened. Infructescences with up to 4 fruits. Fruits sessile, 3-lobed, obtusely triangular from above, heart-shaped in outline, concavely depressed at apex, 57 mm tall, 811 mm diam., dark green when living, densely to sparsely hairy, often becoming glabrous, then the hairs only apical, basal, and in sutures. Seeds semi-globose to ovoid, heart-shaped in outline, sometimes abaxially keeled, 4.55 by 34 by 23.5 mm; testa smooth to rugulate, dark brown.
Distribution Malaysia (fide Airy Shaw, 1975), Borneo, Philippines (Mindanao).
p
= C. glaber; l
= C. angustifolius; n
= C. erycibifolius; q = C.
isabellinus
Habitat & Ecology Primary forests; on brown to black, sandy soil; on hillside (Whitmore, 1973).
Note Cleistanthus glaber is mainly characterised by its glabrousness in branchlets and leaves (the petiole rarely bears some scattered erect hairs) and the greenish yellow leaf colour in the dried state. The fully glabrous C. sumatranus, with which this species could be confused, always dries brownish grey. Cleistanthus bridelifolius is similar in leaf shape, but always has an indumentum. Cleistanthus baramicus from Peninsular Malaysia (?) and Borneo differs from C. glaber by its larger stipules and the brownish colour of the dried leaves.
Cleistanthus isabellinus Elmer, Leafl. Philipp. Bot. 3 (1910) 911; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 40; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 420; Salvosa, Lex. Philipp. Trees (1963) 92; Airy Shaw, Enum. Euphorb. Philipp. Is. (1983) 14; S.Dressler, Blumea 44 (1999) 125, map 1. Lectotype (designated by Dressler 1999: 125): Elmer 12268 (holo NY; iso A 2x, BO, DS, E, F, G 2x, GH, K, L, P, PNH (n.v., photo in A), U, US, WRSL), Philippines, Is. Sibuyan, Prov. Capiz, Magallanes (Mt Giting-Giting).
Tree, up to 20 m high, clear bole at least 7 m high, dbh up to 25 cm. Outer bark dull brown, smooth; inner bark reddish; sapwood thin, whitish; heartwood reddish brown, moderately hard, heavy. Branches finely brownish sericeous when young, later glabrous. Stipules triangular, often scale-like, c. 1.5 by 1 mm. Leaves: petiole terete, rather slender, darker coloured than midrib in dry state, finely appressed hairy, brownish puberulous to sericeous, (5)610 by 11.2 mm; lamina elliptic, rarely slightly oblong, sometimes very slightly obovate, 60115 by 2045 mm, 2.33.8 times as long as wide, chartaceous to (sub)coriaceous, base acute, margin entire, rarely faintly revolute, apex distinctly acuminate, acumen up to 30 mm long above shiny deep green when fresh to shine olive to greenish brown when dry, glabrous with some scattered hairs, beneath glaucous green or yellowish brown when fresh, lighter brown than above when dry, finely appressed lightish to golden puberulous to sericeous (lens!), very young shoots sometimesfulvous hairy; venation: secondary veins in 58(10) pairs, indistinct to subprominent beneath, (sub)prominent above, sometimes tertiary arches present, reticulate areolation. Inflorescences: medium-flowered glomerules (up to c. 10 flowers) axillary on normal-leaved branches; bracts fulvous, ovate triangular, up to 1 by 0.70.8 mm, membranous, brownish puberulous to sericeous outside. Flowers not seen, 36 mm diam. in fruit, odourless. Sepals triangular, 1.53 by 11.5 mm, glabrous to slightly albo-pilose. Infructescences with up to 8 fruits. Fruits subsessile (sometimes pedicel 0.51 mm long), but 24 mm stipitate, stipe glabrous, fruit erect, 11.5 mm diam., obtusely trilobed from above with shallow grooves, depressed oval with concave apical depression in outline, c. 89 mm tall, 810 mm diam., green turning reddish brown, glabrous, sometimes with very few whitish hairs, blackish when dry. Seeds compressed semi-ovoid, c. 4 by 3.5 by 3 mm; testa smooth, dark brown (only 1 seed seen).
Distribution Borneo (Sabah), Philippines (SW Luzon, Sibuyan Island, E Samar), New Guinea (W Irian: Merauke Distr.; Papua: Central Prov.).
q = C.
isabellinus;
l
= C. angustifolius; n
= C. erycibifolius; p
= C. glaber
Habitat & Ecology Humid, primary forests; at altitudes up to 250 m. Reported from gravelly damp soil.
Notes Cleistanthus isabellinus differs from the widespread C. myrianthus merely in the rather acuminate leaf apex and the conspicuous coppery colour of the abaxial leaf indumentum (especially present on the young leaves). Additionally the abaxial lateral veins are far less prominent in C. isabellinus. In all other features, especially in structure of this indumentum, the number of secondary veins and the shape and glabrousness of the capsules, this species is very similar to the rather variable C. myrianthus. Both taxa are characterised by the dense appressed indumentum on the lower leaf surface. It is very likely that further work will allow this species to be included within C. myrianthus. This was already assumed by Jablonsky (1915: 40). More abundant material and field studies should answer this question. I still retain it as separate species which is characterised by the above mentioned character combination.
One collection from North Borneo (SAN (Ambullah) 36010, L) and two from New Guinea (van Royen 4707, Schodde 2953, both L) indicate that this taxon is no Philippine endemic, but although rarely collected more widespread than thought.
Cleistanthus megacarpus C.B.Rob., Philipp. J. Sci., Bot. 6 (1911) 323; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 31; Merr., J. Str. Branch Roy. As. Soc. 86 (1922) 322; Enum. Philipp. Flow. PI. 2 (1923) 420; Holthuis & H.J. Lam, Blumea 5 (1942) 200; Masam., Enum. Phan. Bornear. (1942) 392; Salvosa, Lex. Philipp. Trees (1963) 92; Meijer, Bot. News Bull. Forest Dept., Sabah 7 (1967) 39 in clavi; Airy Shaw, Kew Bull., Add. Ser. 4 (1975) 83; J.A.R. Anderson, Check List Trees Sarawak (1980) 182; Airy Shaw, Kew Bull. 37 (1982) 13; Enum. Euphorb. Philipp. Is. (1983) 14; Whitmore, Tree Fl. Indon., Checkl. Maluku (1989) 39; Tree Fl. Indon., Checkl. Kalimantan 1 (1990) 124; S.Dressler, Blumea 44 (1999) 126, fig. 3, map 4. Lectotype (designated by Dressler 1999: 126): Whitford 1443 (holo K; iso NY), Philippines, Mindoro, Camantigue, Bongabong River.
Shrub to tree, up to 13 m high, stem up to 7 m, dbh up to 26 cm reported. Outer bark dark grey to reddish, smooth; inner bark yellowish red to purplish brown; sapwood white red to light brown. Branches scattered albo-pilose to glabrous. Stipules obliquely narrow triangular, subulate,rather conspicuous, 510 by 13.5 mm, reddish to dark brown, chartaceous to membranous, glabrous, rarely with ciliate margin, margin membranous, conspicuously densely clustered on base of shoots and persistent even without leaves. Leaves: petiole conspicuously long, subterete, conspicuously dark in dry state, glabrous, rarely scattered pilose, 713 by 1.52(2.5) mm; lamina elliptic, often (slightly) obovate, 1035 by 512 cm, 2.23.2 times as long as wide, submembranous to chartaceous, rarely subcoriaceous, base obtuse to acute, margin entire, sometimes slightly revolute, apex acuminate, acumen up to 30 mm long, both sides glabrous, rarely scattered albo-pilose beneath, light green when fresh, often conspicuously glaucescent beneath, greyish brown when dry; venation: secondary veins in 914(17) pairs, prominent beneath,faintly subprominent or lighter coloured above, with rather acute angle of divergence, weak secondary arches, tertiary veins percurrent, reticulate areolation. Inflorescences: few- to medium flowered glomerules (up to 10 flowers) axillary on normal-leaved branches; bracts broadly ovate-triangular, 1.52.5 by 1.52 mm, albo-pilose, dark midvein, margin membranous. Flowers sessile, 45.5(7) mm diam., whitish. Sepals (narrow-)triangular, 23.5 by 12.5 mm, glabrous. Petals whitish, spathulate to rhomboidal ,margin sometimes lobulate, 11.5 by 0.71 mm. Disc glabrous, in staminate flowers and outer one in pistillate flowers saucer-shaped, c. 1.5 mm diam., inner one cup-shaped, c. 1.5 (?) mm tall. Staminal column not seen, pale green to white, free part of filaments up to 0.7 mm long; anthers ovoid, c. 0.7 by 0.4 mm; pistillode cylindrical, with widened blunt apex, glabrous, c. 1 mm tall. Ovary ovoid, c. 1 mm diam., height ?, with a ring of brownish hairs at base (only 1 flower studied); stigmas c. 0.5 mm long. Infructescences suberect. Fruits mostly solitary, occasionally 2- or 3-clustered,a large capsules, sessile, conspicuously stipitate, stipe rather robust, 410 mm long, 24 mm diam.; capsule subglobosely trilobed, obtusely triangular from above, oval in outline, 1823 mm tall, 2025 mm diam., glabrous, rarely tuberculate, light yellow, pinkish green, or brown with a brown calyx, dark brownish when dry. Seeds semi-globosely heart-shaped with ventral groove, 1012 by 910 by 68 mm; testa smooth, dark brown.
Distribution Sumatra (Simaloer Is.), Borneo, Philippines, Sangi and Talaud Is. (Airy Shaw 1975); Morotai (AiryShaw 1982).
l
= C. megacarpus; n = C. rufescens; p = C. spec. A
Habitat & Ecology Reported from humid and primary forests, but also open woods; at low altitude up to 500 m. Reported from rich loamy and wet alluvial soils, and from coral limestone rocks.
Notes This species is somewhat isolated among the taxa studied. Surprisingly there are only a very few collections in flower but many fruiting ones. The main diagnostic characters are: the big, longly and stoutly stipitate capsules (more than 2 cm diam.); the characteristic, long subulate stipules (sometimes densely clustered on the base of shoots and persistent even after the leaves have fallen off) and the long, dark petioles. The leaves reach comparatively large dimensions. The flowers are poorly known due to a lack of material. Apart from the bracts the whole plant is rather glabrous; only the young branches and leaves sometimes have a scattered pilose indumentum.
Cleistanthus myrianthus (Hassk.) Kurz, Prelim. Rep. Forest Pegu, App. A. (1875) cx; Forest Fl. Burma 2 (1877) 370; J.Asiat. Soc. Beng. 42, 2 (1874) 242, in note; Benth. & Hook.f., Gen. PI. 3 (1880) 268 in nota; Fern.-Vill. in Blanco, Fl.Filip., ed. 3, Noviss. App. 4 (1880) 187, t. 353; Gamble, Man. Ind. timb. ed. 1 (1881) 598; Hook.f., Fl. Brit. India 5 (1887) 275; Boerl., Handl. Fl. Ned. Ind. 3 (1900) 271; Brandis, Ind. Trees (1906) 561; C.B.Rob.,Philipp. J. Sci., Bot. 3 (1908) 190; J.J.Sm., Nova Guinea 8 (1910) 231; in Koord. & Valeton, Bijdr. Boomsoort. Java 12 (1910) 297; Koord., Exkurs.-Fl. Java 2 (1912) 484 in clavi; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 37, f. 7 AD; Merr.,Bibl. Enum. Born. Pl. (1921)334, Enum. Philipp. Flow. PI. 2 (1923) 421; Ridl., Fl. Malay Penins. 3 (1924) 194; S.Moore, J. Bot. 63, Suppl. (1925) 93; Gagnep. in Lecomte, Fl. Indo-Chine 5 (1926) 482; Merr., Univ. Calif. Publ. Bot. 15 (1929) 154; Gage, J.Asiat. Soc. Bengal 75, 5 (1936) 515; Masam., Enum. Phan. Bomear. (1942) 392; Holthuis & H.J.Lam, Blumea 5 (1942) 200; K.Heyne, Nutt. PI. Ned.-Ind., ed. 3 (1950) 918; Salvosa, Lex. Philipp. Trees (1963) 92; Backer & Bakh.f., Fl. Java 1 (1964) 474 in clavi; Meijer, Bot. News Bull. Forest Dept., Sabah 7 (1967) 41 in clavi; Airy Shaw, Kew Bull. 26 (1972) 237; Whitmore, Tree Fl. Malaya 2 (1973) 82 in clavi; Airy Shaw, Kew Bull., Add. Ser. 4 (1975) 83; Kew Bull. 31 (1976) 378; Kew Bull. 33 (1978) 43; Kew Bull., Add. Ser. 8 (1980a) 60; Kew Bull. 35 (1980b) 611; Muelleria 4 (1980c) 221; J.A.R.Anderson, Check L. Trees Saraw. (1980) 182; Airy Shaw, Kew Bull. 37 (1982) 13; Enum. Euphorb. Philipp. Is. (1983) 15; Whitmore,Tree Fl. Indon., Checkl. Sumatra (1986) 80; Checkl. Maluku (1989) 39; Checkl. Sulawesi (1989) 46; Checkl. Kalimantan 1 (1990) 124; Hnatiuk, Austral. Fl. Fauna Ser. 11 (1990) 180; A.D.Chapm., Austral. Fl. Fauna Ser. 12 (1991) 740; Pham-hoang Hτ, Cβyco Viκtnam 2, 1 (1992)286; Keίler & Sidiyasa, Trees Balikpapan-Samarinda Area, Tropenbos ser. 7 (1994) 126 in clavi; S.Dressler, Blumea 44 (1999) 129, map 5. Nanopetalum myrianthum Hassk., Verslagen Meded. Afd. Natuurk. Kon. Akad. Wetensch. 4 (1855) 141; Bot. Zeitung (Berlin) 14 (1856) 803; Flora 40 (1857) 534; Hort. Bogor. Descr. (1858) 66; Miq., Fl. Ned. Ind. 1, 2 (1859) 357; Mόll.Arg. in DC., Prodr. 15,2 (1866) 510;Kurz, J. Asiat. Soc. Beng. 42, 2I (1874) 242. Kaluhaburunghos myrianthus (Hassk.) Kuntze, Rev. Gen. PL 2 (1891)607, nom. illeg. Lectotype (designated by Airy Shaw 1980b): Hasskarl (holo BO, not found there by Dressler; iso L, U, possible iso K), W Java, Bantam, S coast.
Cleistanthus cupreus S.Vidal, Revis. Pl. Vasc. Filip. (1886) 235; Airy Shaw, Kew Bull., Add. Ser. IV (1975) PAGE ; Koord., Versl. Minahasa = Meded. Lands Plantentuin 19 (1898) 582; C.B.Rob., Philipp. J. Sci., Bot. 3 (1908) 191. Cleistanthus myrianthus (Hassk.) Kurz subsp. cupreus (S.Vidal) Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 39; Merr., Enum. Philipp. Flow. PI. 2 (1923) 421. Lectotype (Dressler 1999): S. Vidal 560 (holo MA; iso A, AAU, K, L, MA), Philippines, Prov. Rizal, Distr. Morong, Bosoboso.
Cleistanthus apiculatus C.B.Rob., Philipp. J. Sci., Bot. 3 (1908) 189; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 42; Merr., Enum. Philipp. Flow. Pl. 2 (1923)419; Salvosa, Lex. Philipp. Trees (1963) 92; Airy Shaw, Enum. Euphorb. Philipp. Is. (1983) 14. Lectotype (Dressler 1999): Williams 2356 (holo NY; iso GH, K, NY, US), Philippines, Mindanao, Distr. Zamboanga, Sax River.
Cleistanthus pseudocanescens Elmer, Leafl. Philipp. Bot. 3 (1910) 910; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 421; Elmer, Leafl. Philipp. Bot. 4 (1911) 1284. Cleistanthus pseudo-cinereus Elmer ex Airy Shaw, Kew Bull. 33 (1978) 45 (erroneous spelling), nom. inval. Lectotype (designated by Dressler 1999): Elmer 12140 (holo NY; iso A, DS, E, F, G 2x, GH, HBG n.v., K, L, P, U, US, WRSL),Philippines, Is. Sibuyan, Prov. Capiz, Magallanes (Mt Giting-Giting).
Cleistanthus misamisensis C.B.Rob., Philipp. J. Sci., Bot. 6 (1911) 325; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 41; Merr., Enum. Philipp. Flow. PI. 2 (1923) 420; Salvosa, Lex. Philipp. Trees (1963) 92; Airy Shaw, Enum. Euphorb. Philipp. Is. (1983) 15. Lectotype (designated by Dressler 1999): FB (Pray & Cenabre) 15463 (holo US), Philippines, Mindanao, Prov. Misamis, Iligan.
Cleistanthus mindanaensis C.B.Rob., Philipp. J.Sci., Bot. 6 (1911) 324; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 41; Merr.,Enum. Philipp. Flow. PI. 2 (1923) 420; Salvosa, Lex. Philipp. Trees (1963) 92; Airy Shaw, Enum. Euphorb. Philipp. Is. (1983) 15. Lectotype (designated by Dressler 1999): FB (Whitford & Hutchinson) 9474 (holo K; iso L, P, PNH (n.v., photo in A), US), Philippines, Mindanao,Distr. Zamboanga, Port Banga.
Cleistanthus pseudomyrianthus Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 41; Ridl., Fl. Malay Penins. 3 (1924) 194; Gage, J.Asiat. Soc. Bengal 75, 5 (1936) 516; Whitmore, Tree Fl. Malaya 2 (1973) 82. Lectotype (designated by Gage 1936: 517): Wray 2486 (holo K, not seen; iso SING), Peninsular Malaysia, Perak, Larut.
Cleistanthus castus S.Moore, J. Bot. 63, Suppl. (1925) 93; Airy Shaw, Kew Bull. 36 (1981) 281. Type: H.O. Forbes 1622 (iso GH, L, SING), Sumatra, hills S of Goenoeng Trang, Lampongs.
Tree, up to 30 m high, with a rather long and thick stem, clear bole up to 15 m, dbh up to 60 cm. Outer bark greyish, smooth to roughened blotched; inner bark roseous red; sapwood thin, whitish; heartwood rose-red, hard; blaze with a sickly-sweet smell (Airy Shaw, 1980a). Branches finely brownish sericeous when young, later glabrous, quite rigid. Stipules triangular, often scale-like, c. 1.5 by 1 mm. Leaves: petiole terete, rather slender, darker coloured than midrib in dry state, finely appressed hairy, brownish puberulous to sericeous, (5)616 by 12.5(3) mm; lamina elliptic to oblong, rarely lanceolate, sometimes slightly ovate or obovate, 60270 by 1699 mm, 1.84.8 times as long as wide, chartaceous to (sub)coriaceous, base obtuse, roundish to acute, margin entire, rarely faintly revolute, apex acuminate, sometimes acute, acumen up to 45 mm long, above dark green, glossy when fresh to olive to greenish brown when dry, glabrous with some scattered hairs, often rather glossy, sometimes bullate when dry, below greyish green or glaucous when fresh to lighter brown than above when dry, finely appressed lightish to golden puberulous to sericeous beneath (lens!); very young shoots sometimes circinate (fern-like rolled inwards) and fulvous hairy, old leaves die bright yellow; venation: secondary veins in (9)1016(19) pairs, prominent beneath, (sub)prominent above, sometimes tertiary arches present, reticulate areolation. Inflorescences: many-flowered glomerules (more than 10 flowers) axillary on normal-leaved branches, sometimes axillary, spike-like, leafless branches bearing the glomerules, glomerules densely fulvo-pubescent to -sericeous because of bracteal indumentum, and often conspicuously protruding in fruiting state; bracts fulvous, ovate triangular, c. 1 by 0.70.8 mm, membranous, brownish puberulous to sericeous outside. Flowers (sub)sessile, rarely up to 1.5 mm pedicelled (esp. staminate flowers), 36 mm diam., odourless, citrinous to yellow green, often dark brown when dried. Sepals triangular, 1.52.5 by 11.5 mm, glabrous to slightly albo-pilose. Petals spathulate, rhomboid or obtriangular, margin sometimes lobulate,0.61.2 by 0.50.7 mm. Disc glabrous, in staminate flowers and outer one in pistillate flowers saucer-shaped, 1.51.8 mm diam., inner one cupshaped, 11.5 mm tall. Staminal column c. 1 mm long, free part of filaments up to 1.2 mm long; anthers ovoid, 0.51 by 0.30.6 mm; pistillode ovoid, apically turbinate, apex sometimes trifid, glabrous, up to 0.8 by 0.30.4 mm. Ovary semi-globose to ovoid, c. 1 by 11.5 mm, glabrous; stigmas 1.52 mm long, apical quarter bifid, widened at the tip. Infructescences with up to 10 fruits. Fruits subsessile (sometimes pedicel 0.51 mm long), but 25(8) mm stipitate, stipe glabrous, 1.22.5 mm diam.; fruits obtusely trilobed from above with shallow grooves, depressed oval with concave apical depression in outline, 711 mm tall, 79( 14) mm diam., glabrous, sometimes with very sporadic whitish hairs, colouring dull purple when fresh, blackish when dry. Seeds semi-ovoid, adaxially with medial concave depression, apically blunt, 56 by c. 4 by 23 mm; testa smooth, dark brown.
Distribution Burma, Thailand, Andamans, Peninsular Malaysia, Sumatra, Borneo, Philippines, Java, Celebes, Talaud Is., Flores (?, cf. C. myrianthus), Ambon, Morotai, Moluccas (Bacau Is.), New Guinea, Solomon Islands, N Australia (Queensland).
Habitat & Ecology Humid, primary forests; at altitudes up to 800 m. Reported from red or brown or sandy soil, or limestone scree.
Uses Leaves applied to treat asthma (PNH 38375).
Notes This species is very variable with regards to its leaf characters: from narrow elliptic (nearly lanceolate) to broadly elliptic, from small to large, from chartaceous to bullate-coriaceous. Characteristic, however, is the finely appressed indumentum beneath. Additionally, the many-flowered inflorescence glomerules which become conspicuously protruding in fruitand the small, stipitate, glabrous capsules facilitate the recognition of this widespread species. Already in 1874 Kurz recognised the necessity of the new combination(Nanopetalum Hassk. ad Cleistanthus certissime reducendum est.), but without mentioning the epithet. The new combination of Cleistanthus myrianthus was formally made in Kurz (1875: App. A: cx). It is sufficient, as it refers to the basionym. Hence, Chapman's (1991: 740) opinion (This name must be regarded as a new description and not as a new combination as Kurz makes no reference to a basionym. i.e., in his Forest Fl. Burma 1877) is erroneous.
The description above is based mainly on Philippine material where the truly spicate variety (var. spicatus Airy Shaw) was not found.
A var. concinnus (syn. C. pseudocanescens) was described by Airy Shaw based on the manifestly convex leaf, which is glossy on its upper surface. Such collections were made in the Philippines too. However, this character combination badly needs field study to assess its validity for nomenclatura lrecognition.
Cleistanthus apiculatus was based mainly on its narrower and longer acuminate leaves having an apiculate top in comparison with C. myrianthus. Having checked the type-material I am convinced that the former is only a rather narrow-leaved collection of the latter.The apiculate apex is not even a consistent characteristic of all leaves. In all important diagnostic features (e.g., number and prominence of lateral venation, indumentum of leaf and ovary) this material agrees with the very variable C. myrianthus and a separate status cannot be maintained.
Robinson (1911) stated that C. mindanaensis is closely allied to C. isabellinus but differs mainly by larger and thicker, less acuminate leaves with a greater numberof lateral veins (10-13 pairs) and stouter branches. All this clearly matches with C. myrianthus. Only the coppery colour of the undersurface of (especially the young) leaves suggests C. isabellinus, whose specific justification is still somewhat questionable (see there!). Although the young fruits of the type collection show no distinct stipe I am convinced that C. mindanaensis is conspecific with C. myrianthus.
Cleistanthus pedicellatus Hook.f., Fl. Brit. India 5 (1887) 281; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 44; Merr., Enum. Philipp. Flow. PI. 2 (1923) 421; Ridl., Fl. Malay Penins. 3 (1924) 187; Gage, J. Asiat. Soc. Bengal 75, 5 (1936) 498; Whitmore, Tree Fl. Malaya 2 (1973) 80 in clavi; Airy Shaw, Kew Bull., Add. Ser. 4 (1975) 84; Kew Bull., Add. Ser. 8 (1980a) 61; Enum. Euphorb. Philipp. Is. (1983) 15; P.T.Li, Acta Phytotax. Sin. 25 (1987) 139; Whitmore, Tree Fl. Indon., Checkl. Kalimantan 1 (1990) 125; R.I.Milne, Kew Bull. 49 (1994) 452; P.T.Li, Fl. Reipubl. Pop. Sin. 44, 1 (1994) 27; S.Dressler, Blumea 44 (1999) 132, map 6. Kaluhaburunghos pedicellatus (Hook.f.) Kuntze, Rev. Gen. PI. 2 (1891) 607, nom. illeg. Lectotype (designated by Gagge 1936: 499): Curtis 169 (holo K; iso K, SING 8x), Peninsular Malaysia, Penang, Government Hill.
Cleistanthus integer C.B.Rob., Philipp. J.Sci., Bot. 3 (1908) 196; Elmer, Leafl. Philipp. Bot. 3 (1910) 908; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 45, f. 6 EG; Merr., Enum. Philipp. Flow. PI. 2 (1923) 420; Salvosa, Lex. Philipp. Trees (1963) 92; Airy Shaw, Kew Bull. 23 (1969) 65); Enum. Euphorb. Philipp. Is. (1983) 14. Lectotype (designated by Dressler 1999): FB (Aherns collector) 3076 (holo US; iso BO, NY), Philippines, Luzon, Prov. Rizal, Bosoboso.
Cleistanthus quadrifidus C.B.Rob., Philipp. J. Sci., Bot. 3 (1908) 197; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 44; Merr., Enum. Philipp. Flow. PI. 2 (1923) 421; Salvosa, Lex. Philipp. Trees (1963) 92; Airy Shaw, Kew Bull. 23 (1969) 64. Lectotype (designated by Dressler 1999): FB (Whitford & Hutchinson) 9478 (holo US; iso K, NY), Philippines, Mindanao, Distr. Zamboanga, Port Banga.
Cleistanthus dichotomus J.J.Sm. in H.Lorentz, Nova Guinea 8(1912) 786, t. 136; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 43. Type: Gjellerup 142 (iso BO 2x, K, L, U), Niederl. New Guinea [= Papua], N-Kόste bei Biwak Hollandia (Humboldt-Bai).
Cleistanthus pedicellatus Hook.f. f. crassipes Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 44. Type: Wray 2375 (iso K ? n.v., SING), Peninsular Malaysia, Perak.
[Cleistanthus paucinervius Merr. in P.K.Holmgren et al., Index to specimens filed in the New York Bot. Gard. Vasv. PI. Type Herb. (1985) 281, nom.nud., syn. nov. Authentic material: FB (Aloba) 29311 (A, K, NY, US), Philippines, Luzon, Cagayan Prov..]
Cleistanthus monocarpus R.I.Milne, Kew Bull. 49 (1994) 450, f.3. Type: Greagh s.n. (holo K), Borneo, Sabah, Pulau Gaya.
Treelet to tree, up to 16 m high, with a clear bole up to 9 m high, up to 20 cm dbh. Outer bark yellowish grey, peeling in small, shredded plates; inner bark greenish; sapwood greenish white wood hard. Branches glabrous, laxly rebranched, the ultimateones very fine. Stipules ovate triangular to narrow triangular, subulate, small, sometimes tiny, often very inconspicuous, 12.5(6) by 0.51 mm, glabrous. Leaves: petiole subterete, darker coloured than midrib in dry state(blackish), glabrous, sometimes with occasional hairs, (2.5)47 by (0.7) 11.5 mm; lamina elliptic, sometimes ovate, 40145 by 2064 mm, 1.43.3 times as long as wide, (sub)coriaceous, base obtuserotundate to acute or slightly attenuate, margin entire, apex acuminate, acumen 1025 mm long, surfaces greyish brown when dry, glabrous, bright red brown when young; venation: conspicuously prominent on both sides, distinctly reticulate beneath, secondary veins in 48(9) pairs, reticulate areolation. Inflorescences: many-flowered glomerules (up to c. 20 flowers) in the axils of normal leaves, sometimes on short axillary leafless or smaller-leaved shoots; bracts very small, broadly triangular, 0.50.8 by 0.50.8 mm, glabrous, some with occasional hairs or hairy margin, abaxially keeled (midvein), with erose margin. Flowers distinctly pedicelled; pedicel slender, terete, glabrous, (2)513 mm long, 0.20.5 mm diam., those of staminate flowers mostly more slender; flowers 57(8 in fruit) mm diam., creamy white, odourless. Sepals triangular, 23.5 by 1.21.8 mm, glabrous. Petals spathulate to broadly obtriangular, 11.2by 0.61 mm, margin lobulate, midrib darker. Disc in staminate flowers and outer one in pistillate flowers saucer-shaped, 1.52 mm diam., inner one cup-shaped, c. 1 mm tall, covering base of ovary, glabrous, margin irregular. Staminal column 11.5 by c. 0.5 mm, free part of filaments up to 1 mm long; anthers ellipsoid to ovoid, 0.71 by 0.50.8 mm; pistillode ellipsoid, deeply trifid at apex, glabrous, 11.3 by 0.4 0.5 mm. Ovary depressed globose, 11.5 by 1.22 mm, whitish strigose to pilose; stigmas 1.52 mm long, deeply bifid (up to half of their length), widened lobulate to bifidly divided (as quadrifid). Infructescences with up to 9 fruits per glomerule. Fruits distinctly pedicelled, pedicel 713 by 12 mm; fruit a subglobose capsule, depressed, often perfectly round from above, 58 mm tall, 710 mm diam., scattered fulvo-puberulous to later glabrous, hairs especially basal, ascending, yellowish green to reddish to blackish with dark persistent calyx. Seeds semi-globosely broadly heart-shaped, ventrally concavely depressed, dorsally keeled, 3.54.5 by 34 by 23 mm; testa smooth, dark brown.
Distribution China(?; fide Li, 1987), Peninsular Malaysia, Borneo, Philippines, New Guinea.
Habitat & Ecology Primary lowland, also logged over, and secondary forests; up to 300 m. Reported from gravelly, stony, or red clay soil.
Uses Wood locally used for construction or as fuel.
Notes This species is well characterised by slenderly pedicelled flowers and fruits and by subcoriaceous to chartaceous leaves with a distinct prominent venation. It has only a few pairs of secondary veins and is glabrous in all young parts (only very occasionally pilose). Together with the newly recorded C. erycibifolius this species is the only pedicelled one in the Philippines.
Material collected in New Guinea tends to have longer, subulate stipules and denser pilose young fruits.
To me (= Dressler 1999) C. monocarpus represents a weak form of the normal C. pedicellatus as it differs only in having 1 fruit per infructescence and petioles of only 2 mm length. In fact, the paratype (Kahar 5477, L) has petioles of 2.5 mm length.
Cleistanthus pilosus C.B.Rob., Philipp. J. Sci., Bot. 6 (1911) 326; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 33; Merr., Enum. Philipp. Flow. PI. 2 (1923) 421; Salvosa, Lex. Philipp. Trees (1963) 92; Airy Shaw, Enum. Euphorb. Philipp. Is. (1983) 15; S.Dressler, Blumea 44 (1999) 135, map 7. Type: FB (Almagro) 19511 (iso K), Philippines, Basilan.
Treelet to tree, up to 4 m high, dbh 6 cm (only one collection with notes!). Branches long light-coloured-pilose to velutinous when young. Stipules narrow- to oblique-triangular, 1.21.5 by 0.51.5 mm, pilose to glabrous, scaly, inconspicuous. Leaves: petiole subterete, darker coloured than midrib in dry state (blackish), pilose to velutinous, 23 by c. 1 mm; lamina (oblong) elliptic, very rarely slightly obovate, 4090 by 1335 mm, 1.93.3 times as long as wide, chartaceous to subcoriaceous, base subcordate, slightly asymmetric, margin entire, apex shortly acuminate, acumen up to 20 mm long, above glabrous, rarely with occasional hairs on midrib, dull greyish brown when dry, beneath lighter, glabrous except midrib and main veins pilose, sometimes also margin and lower part of blade; venation: indistinct to subprominent above, prominent beneath, secondary veins in 7 or 8 pairs, secondary arches resemble tertiary veins, reticulate areolation. Inflorescences: few-flowered glomerules (not seen) on short spike-like axillary shoots, rarely in the axils of normal leaves, the spike axis pilose; bracts small, triangular, 12 by 0.71 mm, albo-pilose outside. Flowers not seen, sessile, 57 mm diam.(in fruit). Sepals narrow-triangular, c. 3.5 by 11.5 mm, albo-pilose to -sericeous outside. Petals spathulate to roundish, margin often erose, 11.5 by c. 1.5 mm, albo-pilose outside. Inner disc in pistillate flowers up to 1 mm tall. Stigmas 1.52 mm long, apical half of stigmas bifidly divided. Infructescences with 1 (or 2?) fruit(s) per glomerule. Fruits sessile, ob-semiglobose, trilobed from above, compressed oval and concavely depressed at apex in outline, 1011 mm tall, 1012 mm diam., long pilose, becoming glabrescent, but still pilose in grooves and at base. Seeds semi-globose, ventrally and basally concavely depressed, 56 by 4.55 by 34 mm; testa smooth to somewhat striate, light brown.
Distribution Philippines (Basilan and Mindanao).
l
= C. pilosus; n = C. sumatranus
Habitat & Ecology Reported from 60 m altitude.
Uses Used for lumber (FB 27034).
Note This taxon is mainly characterised by the conspicuous pilose indumentum (especially of young twigs, petiole, midrib beneath, inflorescence axis). Furthermore, it has subcordate leaf bases, short spicate inflorescences and its leaves often dry greyish brown. Sessile capsules (which become glabrous), inconspicuous stipules, and albo-pilose petals as well as narrow triangular sepals are other distinctive features. Only a few collections could be affiliated to the species, which shows some morphological similarities with C. bridelifolius.
Cleistanthus robinsonii Elmer, Leafl. Philipp. Bot. 3 (1910) 909; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 52; Merr., Enum. Philipp. Flow. PI. 2 (1923) 422; Salvosa, Lex. Philipp. Trees (1963) 93; Airy Shaw, Enum. Euphorb. Philipp. Is. (1983) 15; S.Dressler, Blumea 44 (1999) 136, map 8. Lectotype (designated by Dressler 1999): Elmer 12540 (holo NY; iso BO, E, F, G, HBG n.v., K, L, P, U, US, WRSL), Philippines, Sibuyan Is., Prov. Capiz, Magallanes (Mt Giting-Giting).
Slender and tall tree, up to 15 m high, trunk reported to c. 60 cm diameter, not straight, with main branches arising from the middle, conspicuously wadded', burly. Bark relatively thin, brown, scaling in plates, reddish beneath the epidermis; wood dingy white on the outside, odourless and tasteless, hard. Branches spreading, numerously rebranched,the ultimate ones very lax and spreading horizontally, appressed brownish pubescent. Stipules scaly, triangular, pubescent, minute, up to 0.5 by 0.5 mm. Leaves: petiole subterete, hardly thicker than midrib and smooth when dry, brownish when dry, fulvously pubescent, 58 by up to 1 mm; lamina linearly oblong, 410 by 12.5 cm, 3.85.2 times as long as wide, submembranous to subcoriaceous, base acute (to round) , margin entire, apex slenderly acuminate, acumen 11.5 cm long, above very scattered albo-pilose, beneath scarcely puberulent, surfaces drying green, glaucescently green beneath; venation: secondary veins in 811 pairs, prominent above and beneath, sometimes with glands (?) in the angle between midrib and secondary veins, areolation indistinct. Inflorescences few-flowered glomerules with up to 6 flowers, axillary on normal-leaved branches, glomerules inconspicuous, with fulvous pubescence; bracts broadly ovate-triangular, minute, c. 0.7 by 0.7 mm, fulvous pubescent. Staminate flowers sessile, 46 mm diam., purple. Sepals triangular, 1.52 by c. 1 mm, glabrous outside. Petals whitish to light yellow, spathulate, 0.51 by 0.30.5 mm. Disc in staminate flowers saucer- to bowl-shaped, c. 2 mm diam., pistillate one not seen. Staminal column 11.5 mm long, free part of filaments up to 1.52 mm long, whitish; anthers ovoid, c. 1 by 0.6 mm, yellowish brown; pistillode conico-ovoid, glabrous, c. 1 by 0.4 mm. Pistillate flowers not seen, style thick, 1.5 mm long (Elmer 1910). Fruits not known.
Distribution Philippines (Palawan, Sibuyan).
l
= C. robinsonii; n = C. venosus
Habitat & Ecology On wooded cliff sand, forested slopes at about 300 m altitude.
Notes This species is still incompletely known. The sterile Loher 4717 collection, determined as C. robinsonii by Airy Shaw, is in my (= Dressler 1999) opinion C. sumatranus. I could trace only one other specimen (Ebalo 634, A) from Palawan, additional to the type collection, of which only some specimens (e.g., NY, WRSL) bear a few flowers. Because of the paucity of material I have merely elaborated the description from the literature and added some minor things from the collections I had at hand. A recollection could possibly clarify its taxonomic situation.
The base of the leaves was erroneously reported to be obtuse (Elmer 1910), but is in fact acute.
Cleistanthus rufescens Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 30; Whitmore, Tree Fl. Malaya 2 (1973) 83; Airy Shaw, Kew Bull. 33 (1978) 49; Kew Bull. 36 (1981) 282; Whitmore, Tree Fl. Indon.,Checkl. Sumatra (1986) 80; S.Dressler, Blumea 44 (1999) 137, map 4. Type: Beccari PS 933 (iso FI-B 2x, n.v., photos seen, L), Sumatra, Padang, Sungei Bulu.
Cleistanthus pseudopodocarpus var. leptopus Airy Shaw, Kew Bull. 27 (1972) 76; Kew Bull., Addit. Ser. 4 (1975) 85. Type: Jacobs 5525A (holo K; iso L), Borneo, Sarawak, 1st Division, Mt. Matang, 25 km W of Kuching.
Cleistanthus pseudopodocarpus Jabl. var. pseudopodocarpus auct. non Jabl.: Airy Shaw, Kew Bull. 27 (1972) 76; Kew Bull., Add. Ser. 4 (1975) 85, pro major parte, cum synon. Airy Shaw 1978: 49).
Cleistanthus pseudopodocarpus auct. non Jabl.: Whitmore, Tree Fl. Malaya 2 (1973) 83 (sec. Airy Shaw 1978: 49).
(Small) tree, up to 10 m high, dbh up to 12 cm. Branches glabrous to brownish tomentose. Stipules (oblique) triangular, 1.52.5by 1.52mm, tomentose to glabrous. Leaves: petiole terete, rather slender, darker coloured than midrib in dry state, scattered pilose to glabrous, (5)714 by 1.52.3 mm; lamina elliptic, sometimes slightly obovate, 95240 by 3590 mm, 2.34 times as long as wide, subcoriaceous to chartaceous, base acute to shortly decurrent, or obtuse, margin entire, apex obtuse sometimes with acuminate tip, acumen up to 15 mm long, glabrous above and beneath, rarely some scattered hairs beneath, especially when young (lens!), greenish olive to dark brown above, sometimes bullate when dry, lighter to greyish brown to light green, glaucous beneath; venation: secondary veins in(7)912 pairs, prominent above and beneath, tertiary veins strongly parallel, reticulate areolation. Inflorescences many-flowered glomerules (more than 10 flowers), axillary on normal-leaved branches, sometimes on axillary, spike-like, leafless shoots, axis then albo- or rufo-strigose or tomentose; bracts ovate triangular, 12.5 by 12.5 mm, albo- to fusco-strigose to -pilose outside. Flowers (sub)sessile, rarely up to 0.5 mm pedicelled, 56(8 in fruit) mm diam., not widely opened. Sepals triangular, 24 by 1.53 mm, appressed brownish tomentose outside. Petals spathulate to semilunate with narrowed base, margin erose, 11.5 by 0.61.2 mm. Staminate disc glabrous, saucer-shaped, c. 2 mm diam. Staminal column 11.7 mm long, free part of filaments up to 1.2mm long; anthers ovoid with sagittate base, 0.70.8 by 0.50.8mm; pistillode ovoid, apically turbinate but blunt on top, light hairs especially at base, 0.81 mm tall, 0.50.7 mm diam. Pistillate flowers not seen; stigmas c. 2 mm long, apical half bifidly divided, widened tip. Infructescences with up to 4 fruits. Fruits subsessile to shortly stalked (pedicel 24 mm long, 11.5 mm diam.) and 24 mm stipitate, stipe densely brownish tomentose, c. 1.5 mm diam.; capsule trilocular, obtusely trilobed from above with shallow grooves, semiglobose with concave apical depression in outline, 810 mm tall, 1013mm diam., pale yellowish green, densely brownish tomentose, endocarp hairy inside. Seeds semi-globose to heart-shaped, adaxially with medial concave depression, dorsally keeled, c. 7 by 56.5 by 4.55 mm; testa smooth to rugose, dark brown.
Distribution Peninsular Malaysia (Airy Shaw 1978), Sumatra, Borneo (Sarawak, Sabah, E Indonesian Borneo),Philippines (Basilan, Mindanao).
n = C. rufescens; l
= C. megacarpus; p = C. spec. A
Habitat & Ecology Lowland primary forests. Reported up to c. 100 m altitude. Growing in sandy loam (one reference).
Notes This species is easily confused with C. myrianthus, but a thorough examination of the leaf underside reveals that it lacks the appressed indumentum of the latter. It does have the conspicuous prominent venation and a sometimes bullate leaf surface when dried, like C. myrianthus, but in the densely brownish tomentose calyx, and the brownish tomentose stipe and capsule it is distinct from C. myrianthus, which has glabrous stipes and fruits.
This species is newly recorded for thePhilippines. Two collections were seen from Basilan Island (BS 15402, Santos 4236) and one from Mindanao (BS 36851). The material clearly matches collections of C. rufescens from Borneo. However, the isotype specimen seen from Sumatra (Beccari PS 933, L) has axillary leafless inflorescences, but matches perfectly otherwise. With this the distinction between C. rufescens and C. bakonensis (see the key in Airy Shaw, 1978: 49) becomes doubtful. Since the material seen of C. rufescens has slightly stronger, a bit bullate, coriaceous leaves, whereas C. bakonensis has chartaceous leaves with a conspicuously prominent venation beneath and the flowers resp. fruits are nearly always on slender leafless 'spikes', I still hesitate to lump them. Clearly, the distinction between 'spicate' and axillary glomerules is not a sharp one. Additionally, indumentum and colour are more variable than thought. The fruits of C. rufescens and C. bakonensis are shortly stipitate and shortly pedicellate sometimes (which was not known yet). The whole complex (C. rufescens, C. bakonensis, C. pyrrhocarpus, C. pseudopodocarpus, etc.) needs critical revision and I think also after Airy Shaw's (1978) key and remarks on the sect. Ferruginosi it is far from understood.
Cleistanthus sumatranus (Miq.) Mόll.Arg. in DC., Prodr. 15, 2 (1866) 504; J.J.Sm. in Koord. & Valeton, Bijdr. Boomsoort. Java 12 (1910) 299; Koord., Exkurs.-Fl. Java 2 (1912) 484 in clavi; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 13, f. 2AE; Merr., Bibl. Enum. Born. PI. (1921) 335; Ridl.,Fl. Malay Penins. 3 (1924) 192; Merr., Philipp. J. Sci. 29 (1926) 381; Masam., Enum. Phan. Bornear. (1942) 393; Backer & Bakh.f., Fl. Java 1 (1964) 474 in clavi; Meijer, Bot. News Bull. Forest Dept., Sabah 7 (1967) 40 in clavi; Airy Shaw, Kew Bull. 26 (1972) 238; Whitmore, Tree Fl. Malaya 2 (1973) 82 in clavi; Airy Shaw, Kew Bull., Add. Ser. 4 (1975) 87; J.A.R. Anderson, Check L. Trees Saraw. (1980) 182; Airy Shaw, Kew Bull. 36 (1981) 282; Kew Bull. 37 (1982) 13; Enum. Euphorb. Philipp. Is. (1983) 16; Whitmore, Tree Fl. Indon.,Checkl. Sumatra (1986) 81; P.T. Li, Acta Phytotax. Sin. 25 (1987) 138; Whitmore, Tree Fl. Indon., Checkl. Maluku (1989) 40; Checkl. Sulawesi (1989) 46; Kiu, Acta Phytotax. Sin. 27 (1989) 456; Whitmore, Tree Fl. Indon., Checkl. Kalimantan 1 (1990) 126; Pham-hoang Ho, Cayco Vietnam 2,1 (1992) 288; Siemonsma & Kasem Piluek (eds.), PROSEA 8 (1993) 286; P.T. Li, Fl. Reipubl. Pop. Sin. 44, 1 (1994) 25, t. 7, f. 48; S.Dressler, Blumea 44 (1999) 139, map 7. Leiopyxis sumatrana Miq., Fl. Ned. Ind., Eerste bijv. (1861) 446. Kaluhaburunghos sumatranus (Miq.) Kuntze, Rev. Gen. PI. 2 (1891) 607, nom. illeg. Type: Teijsmann s.n. (holo U; iso Bf, BO, G-DC (microfiche seen), GH n.v., K, L), S Sumatra, Prov. Lampong, in monte Gunung Batin, prope Tega-nennin.
Cleistanthus blancoi S.Vidal, Revis. PI. Vasc. Filip. (1886) 234, nom. illeg., non C. blancoi Rolfe, J. Linn. Soc., Bot. 21 (1884) 315. Cleistanthus vidalii C.B.Rob.,Philipp. J. Sci., Bot. 3 (1908) 193; C.B.Rob., Philipp. J. Sci., Bot. 6 (1911) 327; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 15; Merr., Enum. Philipp. Flow. PI. 2 (1923) 422; Salvosa, Lex. Philipp. Trees (1963) 93; Airy Shaw, Enum. Euphorb. Philipp. Is. (1983) 16. Kaluhaburunghos blancoi (S.Vidal) Kuntze, Revis. Gen. PI. (1891) 607, nom. illeg. Lectotype (designated by Dressler 1999): Vidal y Soler 559 (holo MA; iso AAU, K, L, MA), Philippines, Luzon, Prov. Tarlac, Moriones.
Cleistanthus heterophyllus Hook.f., Fl. Brit. India 5 (1887)276; Hallier f., Meded. 's Rijks Herb. 1 (1910) 7; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 14; Merr., Bibl. Enum. Born. PI. (1921) 334; Ridl., Fl. Malay Penins. 3 (1924) 190; Gage, J.Asiat. Soc. Bengal75,5 (1936)505; Masam., Enum. Phan. Bornear. (1942) 392. Lectotype (designated by Dressler 1999): Maingay KD 1372 (holo K; iso K, L), Malacca.
Cleistanthus laevis Hook.f., Fl. Brit. Ind. 5 (1887) 277; C.B.Rob., Philipp. J. Sci., Bot. 3 (1908) 193; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 13. Type: Marton 109 (holo K), Singapore, jungle behind the Botanical Gardens.
Cleistanthus minahassae Koord., Meded. Lands Plantentuin 19 (1898) 625; J.J.Sm. in Koord. & Valeton, Bijdr. Boomsoort. Java 12 (1910) 301, in obs.; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 52. Lectotype (designated by Dressler 1999): Koorders 16920 (holo L; iso BO), Celebes, Prov. Minahassa, Menado.
Cleistanthus laevigatus Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 12; Merr., Bibl. Enum. Born. PI. (1921) 334; Masam., Enum. Phan. Bornear. (1942) 392; Meijer, Bot. News Bull. Forest Dept., Sabah 7 (1967) 40 in clavi; Airy Shaw, Kew Bull. 21 (1968) 363, in obs. Lectotype (designated by Dressler 1999): Beccari PB 3882 (holo K; iso FI-B, photo seen), Borneo, Sarawak.
Cleistanthus oligophlebius Merr., Philipp. J. Sci., Bot. 13 (1918) 80; Merr., Bibl. Enum. Born. PI. (1921) 335; Univ. Calif. Publ. Bot. 15 (1929) 154; Masam., Enum. Phan. Bornear. (1942) 393. Lectotype (designated by Dressler 1999): Villamil 399 (erroneously in protologue sub no. 339?) (holo K), British North Borneo, Marutai watershed nr Tawau.
Cleistanthus saichikii Merr., Philipp. J. Sci. 23 (1923)248; Gagnep. in Lecomte, Fl. Indo-Chine 5 (1926) 499, f. 64.57; Merr. & Chun, Sunyatsenia 1 (1930) 64; Chun & Chang (eds.), Fl. Hainanica 2 (1965)141; Anonymous, Iconogr. Corm. Sin. 2 (1972) 592, f. 2914. Type: Canton Christian College (McClure) 9148 (iso A 2x, CAS, E, K, NY 2x, P), China, Hainan, Fan Ta.
Paracleisthus subgracilis Gagnep., Bull. Soc. Bot. France 70 (1923) 500; in Lecomte, Fl. IndoChine 5 (1926) 500. Syntypes: Harmand 81 (syn P?, n.v.), Cochinchine, Chaudoc; Lefevre 252 (syn P?, n.v.), Cochinchine, Point A; Magnien, Gourgand etc. s.n. (syn P?, n.v.), Cambodge, Pnom-penh; Pierre 6300 (E, K: s.n., P?, n.v.), Cambodge, Phu-quoc; Poilane 165 (E, K, P?, n.v.), Cochinchine, Sang-dinh.; Poilane 172 (syn P?, n.v.), Cochinchine, Gia-ray; Thorel s.n. (syn P?, n.v.), Laos merid.,Bassac; Thorel s.n. (BR, K, P?, n.v.), Cochinchine.
Cleistanthus euphlebius Merr.,Pap. Michigan Acad. Sci. 24 (1939) 78; Airy Shaw, Kew Bull. 21 (1968) 363, in syn. Type: Rahmat si Toroes 818 (iso A n.v., E, MICH n.v., NY n.v., SING), Sumatra, E Coast, Asahan, Silo Maradja, near Taloen Djoring.
Tree, up to 18 m high, with a clear bole up to 12 m tall, dbh up to 35 cm. Outer bark brownish to pale grey, finely cracked; inner bark thin, white to pale brown; sapwood reddish, hard. Branches glabrous, rarely scattered albo-pilose when young. Stipules (oblique) triangular, 11.5 by 0.81.5 mm, glabrous, rarely with some hairs. Leaves: petiole subterete, darker coloured than midrib in dry state (blackish), glabrous, rarely some scattered hairs adaxialyl, 26(7) by 0.71.2(1.5) mm; lamina (narrowly) ovate to (ovately) elliptic, rarely lanceolate, 27150 by 1155 mm, 2.24.3 times as long as wide, chartaceous to subcoriaceous, base acute to roundish, rarely obtuse, margin entire, apex (shortly) acuminate to acute, acumen 1525 mm long, glabrous, brownish olive and often conspicuously shiny above, brownish grey beneath when dry, hardly glaucescent beneath, but paler greyish when dry; venation: obscure above, faintly prominent beneath, secondary veins in (4) 58 pairs, often acute angles of divergence, long running towards apex before looping, sometimes tertiary arches, irregular reticulate areolation. Inflorescences: few-flowered glomerules (up to c. 7 flowers) in the axils of very small leaves or on leafless spike-like shoots, glomerules resemble 'pearls-on-a-string'; bracts small, triangular, c. 1.5 by 11.5 mm, glabrous to albo-pilose, with sericeous margin. Flowers sessile, 45 mm diam., brownish yellow with red. Sepals (narrow-)triangular, 22.5 by 11.5 mm, glabrous with occasional hairs at apex to albo-pilose outside. Petals narrowly spathulate, 0.71.2 by 0.30.4 mm, margin slightly lobulate. Disc in staminate flowers and outer one in pistillate flowers saucer-shaped, c. 1.5 mm diam., inner one cup-shaped, 0.71.2 mm tall, covering base of ovary, glabrous. Staminal column 11.5 mm long, free part of filaments up to 1 mm long; anthers ellipsoid, 0.30.5 by 0.20.4 mm; pistillode ovoid to ellipsoid, glabrous with some hairs at base, 0.60.8 by c. 0.3 mm. Ovary globose, 11.2 by 11.2 mm, densely albo-strigose; stigmas c. 11.3 mm long, basally united, apical thirds of bifidly divided, tips widened. Infructescences with 1 or 2 fruits per glomerule. Fruits sessile, rarely minutely stipitate (c. 1 mm), obtusely triangular and deeply 3-lobed from above or tricoccous-subglobose, compressed oval and concavely depressed at apex in outline, 79(13) mm tall, 810(13) mm diam., red when ripe, glabrous, basally and in sutures (grooves) with scattered pilose hairs. Seeds ovoid with blunt apex to heart-shaped, ventrally with median hilum, dorsally slightly keeled, 56 by 44.5 by 34 mm; testa smooth, slightly lineate with occasional inconspicuous warts, brown.
Distribution China:Guangdong (fide Kiu, 1989), Hainan, Indochina,Thailand, Peninsular Malaysia, Sumatra, Borneo, Philippines, Java, Celebes, Bali, Ambon.
n = C. sumatranus;
l
= C. pilosus
Habitat & Ecology Dry to alluvial primary forests, also in secondary forests; up to 700 m altitude. Reported from rocky (granitic) alluvial soil and limestone.
Notes This species has a rather wide distribution in SE Asia. Nevertheless it is not exceedingly variable and hence easily recognizable, especially by its characteristic smaller leaved to leafless axillary shoots bearing the flower glomerules (pearls-on-a-string' appearance). It is also characterised by its glabrous, chartaceous to subcoriaceous leaves, which are often glossy above and dry greyish brown, by its few secondary vein pairs, which show a conspicuously acute angle of divergence and run widely towards the apex before looping. It might be confused with C. venosus, which is larger in most parts. The bigger leaves of the latter are more oblong and have more secondary veins which do not run as far towards the apex and which are conspicuously prominent whereas the lateral veins in C. sumatranus are remarkably obscure. The leaves of the former dry conspicuously yellowish or greyish green. The two species are doubtlessly very closely related.
Cleistanthus gracilis from Peninsular Malaysia and Borneo is supposed to differ from C. sumatranus merely by smaller leaves and their distinct acumen. These characters should be carefully compared. Both taxa might well prove to be conspecific.
Cleistanthus venosus C.B.Rob.,Philipp. J. Sci., Bot. 3 (1908) 192; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 12; Merr., Bibl. Enum. Born. Pl. (1921) 335; Enum. Philipp. Flow. PI. 2 (1923) 422; Masam., Enum. Phan. Bornear. (1942) 393; Salvosa, Lex. Philipp. Trees (1963)93; Airy Shaw, Kew Bull., Add. Ser. 4 (1975) 87; Kew Bull. 36 (1981) 282; Enum. Euphorb.Philipp. Is. (1983) 16; Whitmore,Tree Fl. Indon., Checkl. Sumatra (1986) 81; Checkl. Kalimantan 1 (1990) 126; S.Dressler, Blumea 44 (1999) 341, map 8. Lectotype (designated by Dressler 1999): Williams 2187 (holo NY; iso GH, K, NY, US), Philippines, Mindanao, Distr. Zamboanga, Sax River.
Cleistanthus elmeri Merr., Univ. Calif. Publ. Bot. 15 (1929) 154; Masam., Enum. Phan. Bornear. (1942) 392; Meijer, Bot. News Bull. Forest Dept., Sabah 7 (1967) 40 in clavi; Bot. Bull. Herb. Forest Dept., Sabah 10 (1968) 232. Lectotype (designated by Dressler 1999): Elmer 21694 (holo PNHf ?, replaced by L; iso A, B, BO, BR, DS, K, L, M, NY, P, PH, SING, U, US), British N Borneo, Elphinstone Prov., near Tawao.
Treelet to tree, up to 12 m high, with a clear bole up to 9 m high, girth up to 1 m. Outer bark light green to pale grey, smooth; inner bark pale to brownish green; sapwood white to whitish brown. Branches very young scattered pilose, later glabrous. Stipules triangular, 24 by c. 1 mm, glabrous, occasionally some hairs at apex. Leaves: petiole subterete, darker coloured than midrib in dry state (blackish), glabrous, 59(10) by 12.5 mm; lamina elliptic, very rarely slightly obovate, 75270by 20100 mm, 2.64.3 times as long as wide, chartaceous, base acute to roundish, slightly asymmetric, rarely slightly emarginate, margin entire, apex acuminate, acumen 1030 mm long, glabrous, greenish to yellowish when dry, hardly glaucescent beneath, but pale greenish when dry; venation: conspicuously prominent beneath, secondary veins in 58(10) pairs, secondary arches resemble tertiary veins, irregular reticulate areolation. Inflorescences: few-flowered glomerules (mostly 3 or 4 flowers, rarely up to c. 10 flowers) in the axils of very small, lanceolate leaves which are conspicuously hairy beneath, these on spike-like branches in the axils of normal leaves with the spike axis sometimes scattered pilose; bracts small, triangular, 1.52.3 by 1.22 mm, albo-pilose to -strigose outside, abaxially keeled (midvein), with erose margin. Flowers sessile, staminate ones slender, 47 mm diam. Sepals narrow-triangular, 23.5 by 11.5 mm, appressed albo-strigose outside. Petals spathulate to obovate, margin often erose, (1)1.52 by (0.3)11.5 mm. Disc in staminate flowers saucer-shaped, 1.52.5 mm diam., outer one in pistillate flowers saucer-shaped, 23 mm diam., inner one cup-shaped, 11.5(2) mm tall, covering base of ovary, glabrous to weakly pilose outside. Staminal column 1.21.5 by c. 0.5 mm, free part of filaments up to 1.2 mm long; anthers ovoid, 0.71.2 by 0.50.8 mm; pistillode ovoid to globose, apex blunt, glabrous, 11.3 by c. 0.6 mm. Ovary globose, 1.42.2 by 1.52.2, densely albo-strigose; stigmass c. 2 mm long, apical third bifidly divided, papillose, widened. Infructescences with 1 or 2 fruits per glomerule. Fruits (sub)sessile (to 1 mm stalked), obtusely triangular and deeply 3-lobed from above, compressed oval and concavely depressed at apex in outline, 911 mm tall, 913 mm diam., reddish green, scattered pilose, hairs only apical, basal, and in sutures (grooves). Seeds broadly bean-shaped, ventrally and basally concavely depressed, dorsally keeled, c. 7 by 4 by 4.5 mm; testa smooth, slightly lineate with occasional inconspicuous warts, dark brown.
Distribution Peninsular Malaysia, Sumatra (Simaloer Is.), Anambas Is., Borneo, Philippines (Mindanao).
n = C. venosus;
l
= C. robinsonii
Habitat & Ecology Primary lowland forests; up to 600 m altitude; on brown to black, sandy soil.
Notes The smaller-leaved, spike-like flower bearing branches generally characterise C. venosus together with the conspicuous prominence of venation of higher order (3° and more) on the lower leaf surface. Moreover, the chartaceous leaves are glabrous with only scattered hairs beneath. They often dry greenish. The narrow triangular sepals show a strong indumentum, the deeply lobed, sessile capsules are pilose but become glabrous later. From C. everettii it differs by the glabrous petiole and the fewer-flowered glomerules. The leaves often reach a larger size too. Field studies have to prove whether these characters are consistent and both taxa are really distinct. BS (McGregor) 18614 seems to represent a collection intergrading between both.
From C. sumatranus it is distinguished by the larger stipules, the colour of the dried leaves and by shape ofthe flower-bearing branches.
Airy Shaw (1975) states: "Closely related to C. winkleri, with which it shares rather large flowers and long narrow sepals, but differing in that the sepals and petals are externally long pilose". In my opinion the distinction between these taxa should be re-examined when better material is available.
Cleistanthus vestitus Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 32; Merr., Bibl. Enum. Born. PI. (1921) 335; Gage, J. Asiat. Soc. Bengal 75, 5 (1936) 514; Masam., Enum. Phan. Bornear. (1942) 393; Whitmore, Tree Fl. Malaya 2 (1973) 83 in clavi; Airy Shaw, Kew Bull., Add. Ser. 4 (1975) 87; J.A.R.Anderson, Check List Trees Sarawak (1980) 182; Airy Shaw, Kew Bull. 36 (1981) 282; Whitmore, Tree Fl. Indon., Checkl. Sumatra (1986) 81; Checkl. Kalimantan 1 (1990) 126; S.Dressler, Blumea 44 (1999) 143, map 3. Lectotype (designated by Dressler 1999): Teijsmann HB 11331 (holo L; iso BO), Borneo, Sungei Landak.
Cleistanthus vestitus Jabl. f. perakensis Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 32. Cleistanthus perakensis (Jabl.) Gage ex Ridl., Fl. Mal. Penins. 3 (1924) 195 (synonymy based on Gage, J. Asiat. Soc. Bengal 75:, 1936: 514; although Gage does not explicitly mention this species in the synonymy, he cites both syntypes of C. perakensis under C. vestitus, which has to be considered as a synonymisation). Syntypes: Scortechini 1822 & 1823 (K), Peninsular Malaysia, Perak.
Cleistanthus barrosii Merr., Philipp. J. Sci. 20 (1922) 400; Merr., Enum. Philipp. Flow. PI. 2 (1923) 419; Salvosa, Lex. Philipp. Trees (1963) 92; Airy Shaw, Enum. Euphorb. Philipp. Is. (1983) 14. Lectotype (designated by Dressler 1999): FB (Barros) 26070 (holo US), Philippines, Luzon, Isabella Prov., Ilagan.
Cleistanthus sp.: Merr., Univ. Calif. Publ. Bot. 15 (1929) 155 (based on Elmer 21567).
Bridelia cinnamomea auct. non Hook.f., Fl. Brit. India 5 (1887) 273, p.p. (cf. Gage, 1936: 514).
Shrub to small tree, up to 6 m high. Outer bark dark brown, smooth, thin; sapwood yellowish. Branches fulvo-pubescent to -pilose when young. Stipules (narrow) triangular, 24 by 11.5 mm, pubescent to glabrous. Leaves: petiole (sub-)terete, darker coloured than midrib in dry state, light brownish pubescent to pilose, later glabrous, 49 by 12 mm; lamina elliptic, often slightly ovate, sometimes lanceolate, 520 by 28 cm, 23.5 times as long as wide, chartaceous, base obtuse, roundish to acute, rarely attenuate, margin entire, apex shortly acuminate to acute, acumen up to 20 mm long,scattered albo-to fulvo-pilose above, later glabrescent, rusty pubescent to pilose beneath, becoming subglabrous, indumentum at base and main veins usually denser; surfaces often conspicuously dark reddish brown when dry; venation: secondary veins in 69(10) pairs, prominent beneath, (sub)prominent above, with very weak secondary arches, sometimes arches tertiary, perfect reticulate areolation. Inflorescences: conspicuously reddish-woolly, many-flowered glomerules axillary on normal-leaved branches, sometimes axillary, spike-like, leafless branches bearing the glomerules; bracts fulvous, ovate or broadly triangular, c. 2 by 1.5 mm, membranous, often conspicuously brownish strigose to pubescent outside, sometimes glabrous with hairy midvein, with erose margin. Flowers sessile, 34(5) mm diam., brown. Sepals triangular, 1.52.5 by 0.71.5(2) mm, brownish pubescent. Petals whitish, spathulate to rhomboidal, margin lobulate, 0.61 by 0.50.9 mm. Disc glabrous, in staminate flowers and outer one in pistillate flowers saucer-shaped, 1.52 mm diam., inner one cup-shaped, c. 0.5 mm tall. Staminal column up to 1.5 mm long, tree part of filaments up to 1 mm long; anthers ovoid to ellipsoid, 0.50.7 by 0.30.4 mm; pistillode cylindrical, with widened blunt apex, glabrous, up to 1.2 by 0.40.6 mm diam. Ovary (semi)globose, c. 1 by 1.5, densely brownish pubescent; styles c. 1 mm long, apical quarter bifidly divided, widened lobulate tip. Infructescences with up to 5 fruits seen. Fruits sessile, but 24(5) mm stipitate, stipe rather slender, 12 mm diam.; capsule obtusely triangular from above, shallowly tri-lobed, more or less rectangular to broadly ovate in outline, 67 79 mm, light brown or brownish grey, scattered pilose to glabrous, hairs mainly in grooves and at base, brownish when dry. Seeds heart-shaped in outline, abaxially keeled, c. 4 by 3 by 2.53 mm; testa smooth, brown.
Distribution Peninsular Malaysia, Sumatra, Borneo, Philippines (Luzon).
n = C. vestitus;
l
= C. decurrens
Habitat & Ecology Reported from disturbed or secondary forests, but infrequently collected; on low altitudes up to 300 m. Reported from sandy, clayey soil.
Notes This species has its morphologically closest relative in C. decurrens, but can be recognised by the conspicuously reddish tomentose, many flowered glomerules and non decurrent leaf base.
Cleistanthus vestitus is sometimes confused with the very variable C. myrianthus because of its leaf shape and the stipitate capsules. The latter differs, however, sufficiently in its appressed abaxial leaf-indumentum. Additionally it has mostly glabrous capsules. Despite the fact that another Philippine species was named C. bridelifolius I think the leaves of C. vestitus resemble much more closely those of some Bridelia species (see under C. bridelifolius).
The typical C. barrosii described as aPhilippine endemic never has spicate inflorescences, a feature which sometimes characterises C. vestitus (and is noted in its protologue). However, this is not consistent among the material studied of the latter from Borneo and Sumatra. I therefore confirm Airy Shaw's synonymisation.
Cleistanthus spec. A
S.Dressler, Blumea 44 (1999) 144, map 4.
p = C. spec. A; l
= C. megacarpus; n = C. rufescens
I (= Dressler 1999) have seen three collections, all from Palawan, which in my opinion do not match any otherof the species known fromthePhilippines: Ridsdale 1008 (A, BO, K, L 2x), Soejarto et al. 6833 (A, CHI n.v., F n.v., K, L, PNH n.v.), Soejarto et al. 8412 (A n.v., CHI n.v., F, L, PNH n.v.). I also could not relate them to a Bornean species. They might belong to a yet undescribed taxon, which I did not venture to describe, as the material is rather poor in the generative phase (no flowers, only dehisced capsule bases present).
This taxon is characterisedby the following features; Tree up to 25 m high, dbh 20 cm with dry, brown erect inflorescences, being glabrous on branches, petioles, leaves, sepals (only young shoot tips are covered by a brownish indumentum). Stipules triangular, scaly, tiny, c. 0.7 by 0.5 mm. Leaves with 310 mm long petioles; blade lanceolate, elliptic to slightly obovate, rather small, 210 by 0.84 cm, coriaceous, base obtuse to acute, margin somewhat revolute, apex (shortly) acuminate, glossy above; secondary venation when dried conspicuously raised, 810 pairs of secondary veins. Flowers in few-flowered glomerules on normal leaved branches; bracts broadly ovate-triangular with brownish pubescent indumentum. Sepals triangular, c. 2 by 1 mm, glabrous. Capsules sessile, not stipitate, glabrous (at least basally), c. 56 mm tall.
All these collections originate from ultrabasic slopes (collector's statement on two labels; the third, Soejarto et al. 6833, was foundin forest of low stature on a thin soil layer). The plant appears to be common in the area and withstands rather dry conditions.
Cleistanthus orgyalis (Blanco) Merr., Rev. Blancos Fl. Filip. (1905) 75, nom. rej. prop. (S.Dressler & Petra Hoffmann, Taxon 47, 1998: 753); C.B.Rob., Philipp. J. Sci., Bot. 3 (1908) 189; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 42; Merr., Sp. Blancoan. (1918) 220; Merr., Enum. Philipp. Flow. PI. 2 (1923) 421; Salvosa, Lex. Philipp. Trees (1963) 92; Airy Shaw, Enum. Euphorb. Philipp. Is. (1983) 15; S.Dressler, Blumea 44 (1999) 145 (doubtful species). = Gluta orgyalis Blanco, Fl. Filip, ed. 2 (1845) 451. Type: not designated.
Cleistanthus blancoi Rolfe, J. Linn. Soc., Bot. 21 (1884) 315; Merr., Rev. Blancos Fl. Filip. (1905) 75, in syn.; C.B.Rob., Philipp. J. Sci., Bot. 3 (1908) 193; Hallier, Meded. 's Rijks Herb. 1 (1910) 7; Jabl. in Engl., Pflanzenr. IV.147.viii (1915) 13; Merr., Enum. Philipp. Flow. PI. 2 (1923) 419; Salvosa, Lex. Philipp. Trees (1963) 92; Airy Shaw, Enum.Euphorb. Philipp. Is. (1983) 14. = Cleistanthus ferrugineus auct. non Mόll.Arg.: Fern.-Vill. in Blanco, Fl. Filip., ed. 3, Noviss. App., 4 (1880) 187. Type: Fernandez-Villar in Blanco, Fl. Filip., ed. 3, Noviss. App. (1880) t. 353.
Note: The true identity of the Blanco species Gluta orgyalis always remained obscure, as there obviously is no herbarium material extant. An analytical plate in Fernandez Villar (1880: t. 353) was drawn after Blanco's death and therefore does not represent original material. This plate, however, is the type of C. blancoi Rolfe. Unfortunately, it is ambiguous as the depicted parts of the plant do not sufficiently show diagnostic characters. In the article by Dressler & Hoffmann (1998) it is proposed to reject the comparatively early combination C. orgyalis in order to prevent possible undesirable nomenclatural changes. For a detailed discussion see the cited paper.
