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1a. |
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1b. |
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2a. |
Branchlets, petioles and leaves completely glabrous |
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2b. |
Branchlet, petioles and leaves on midrib beneath sparsely puberulous |
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3a. |
Leaves 6.5–18 cm long, smooth or young ones irregularly lowly granular beneath |
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3b. |
Leaves 13–32 cm long, strongly white-granular beneath on nervation only |
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4a. |
Leaf: nerves 5–10 pairs, drying approximately the same shade of grey-yellow-green as the lamina. Inflorescences: glomerules more or less evenly set along the rachis |
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4b. |
Leaf: nerves 9–12 pairs, drying often beneath lighter green than the lamina. Inflorescences: glomerules apically less spaced than at base |
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5a. |
Leaf: nerves 5–8 pairs; texture slightly leathery, drying dull greyish green to dark green; dots densely set, greyish |
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5b. |
Leaf: nerves 7–10 pairs; texture papery, drying shiny yellowish green or dark green; dots indistinct |
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6a. |
Stipules (narrowly) ovate, sometimes slightly oblique, or falcate; caducous, but still present in young stages. Leaf: base cuneate to rounded; nerves 6–13 pairs |
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6b. |
Stipules falcate, persistent. Leaf: base emarginate-truncate; nerves 14–23 pairs |
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7a. |
Branchlets sparsely tomentose to subglabrous. Leaves only sparsely tomentose to subglabrous beneath; drying yellowish green on both sides |
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7b. |
Branchlets tomentose. Leaf: at least (sparsely) tomentose beneath; drying dark brownish to greenish brownish above, brown beneath |
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8a. |
Leaves 5.5–13.5 cm long; dots irregularly densely set, greyish, fading. Inflorescences clustered up to 5 together, 7–17 mm long, 1.5–2 mm diam.; glomerules globose. — North Borneo, Mt Kinabalu and environments, above 1000 m |
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8b. |
Leaves 10–19 cm long; dots absent. Inflorescences clustered up to 15 together, 9–35 mm long, 0.5–1.5 mm diam.; glomerules ellipsoid. — Sarawak, along or partly in rivers or at tidal influence zones |
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9a. |
Stipules (narrowly) ovate, present in young stages. Leaf: marginal glands few, small, indistinct; texture rather papery; (sparsely) hirsute on both sides. Inflorescence rachis 0.3–0.5 mm diam.; glomerules 0.8–1.2 by 0.5–0.8 mm |
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9b. |
Stipules falcate, often early caducous. Leaf: marginal glands many, large, distinct; texture rather leathery; only (sparsely) tomentose beneath. Inflorescence rachis 0.8–1.5 mm diam.; glomerules 1.5–2.5 by 1.5–2.5 mm |
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10a. |
Leaves 11–25 by 5–11 cm. Inflorescences 10–20 clustered together, 11–47 mm long |
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10b. |
Leaves 5.5–13 by 2–4.5 cm. Inflorescences 2–5 clustered together, 7–12 mm long |
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11a. |
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11b. |
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12a. |
Petiole and leaf at least beneath puberulous. Inflorescences continuous, glomerules uninterrupted, indistinct |
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12b. |
Petiole and leaf completely glabrous. Inflorescences continuous, glomerules slightly interrupted, distinct |
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13a. |
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13b. |
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46c. A. octandra var. malesiana |
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14a. |
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14b. |
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15a. |
Leaves glabrous or only beneath sparsely hairy; nerves slightly prominent to flat above |
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15b. |
Leaves: above at least on midrib (sparsely) hirsute, beneath (sparsely) hirsute; nerves more or less sunken above |
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16a. |
Stipules 6–20 by 11–37 mm, often long persistent. Petiole: upper pulvinus swollen, 5–11 by 3–6 mm. Leaves 14–47 cm long; base cordate; nerves raised beneath, not fading towards the margin |
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16b. |
Stipules 4–8 by 11–18 mm, present in young stages. Petiole: upper pulvinus distinct, but not extremely so, 2–6 by 1.5–2.5 mm. Leaves 8–25 cm long; base emarginate to rounded; nerves prominent beneath, fading towards the margin |
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17a. |
Nervation strongly sunken above, giving the leaf a bullate appearance |
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17b. |
Only nerves sunken, sometimes near margin and apex some veins sunken, otherwise leaves flat, smooth above |
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18a. |
Leaves 9–20 cm long |
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18b. |
Leaves 21–29 cm long |
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19a. |
Branchlets and petiole very sparsely puberulous to glabrous; leaves completely glabrous or only very sparsely puberulous on midrib beneath near base. Inflorescences: rachis (sub)glabrous |
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19b. |
Branchlets and petiole (sparsely) puberulous; leaves at least on midrib beneath (sparsely) puberulous to tomentose. Inflorescences: rachis (sparsely) puberulous to tomentose |
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20a. |
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20b. |
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21a. |
Leaf: apex acuminate to cuspidate; texture papery or leathery; drying greyish to yellowish green on both sides, sometimes greenish or brownish above, brown beneath but then apex acuminate and texture leathery; nerves 7–12 pairs. Glomerules interrupted at base, or if only minutely spaced at base, then inflorescences more then 2 mm diam.; inflorescences drying yellowish or brownish |
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21b. |
Leaf: apex cuspidate; texture papery; drying greenish or bluish brown above, often blackening, (light) brown beneath; nerves 4–6 pairs. Glomerules only at base slightly spaced and inflorescences 1–1.3 mm diam.; inflorescences drying dark brownish |
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22a. |
Leaf: texture papery, dull; drying greyish green to yellowish green on both sides; dots absent or indistinct. Glomerules 0.8–2 mm long. Flowers drying yellowish; stamens not to slightly exserted, shorter than to slightly longer than sepals |
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22b. |
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23a. |
Branchlets, petiole, and leaf on midrib beneath very sparsely puberulous. Inflorescences 5–22 mm long, 1–3.5 mm diam.; glomerules ellipsoid, at base interrupted, distinct, apically continuous, indistinct |
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23b. |
Branchlets, petiole, and leaf completely glabrous. Inflorescences 14–41 mm long, 0.8–1.2 mm diam.; glomerules subglobose, strongly interrupted at base, at apex less, but distinct throughout |
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24a. |
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24b. |
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25a. |
Inflorescences 1–6 clustered together; glomerules subglobose to ellipsoid, distinct to indistinct, (slightly) interrupted at base, decreasingly spaced towards the apex |
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25b. |
Inflorescences 4–14 clustered together; glomerules ellipsoid, distinct, spaced rather evenly interruptedly along the rachis. — Inflorescences 9–35 mm long, 0.5–1.5 diam.; glomerules 1–1.5 by 0.5–0.8 mm. Sarawak, along or partly in rivers or at tidal zones |
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26a. |
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26b. |
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27a. |
Branchlets and leaves tomentose. Leaf: nerves 10–14 pairs. Inflorescences 12–32 mm long, tomentose; apical head 5–9 mm diameter |
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27b. |
Branchlets and leaves sparsely tomentose. Leaf: nerves 8–10 pairs. Inflorescences 5–22 mm long, (sparsely) tomentose; apical head 2–3.5 mm diameter |
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28a. |
Leaves: nerves 4–10 pairs |
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28b. |
Leaves: nerves 10–17 pairs |
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29a. |
Leaf: apex acuminate to cuspidate; texture (rather) leathery; dots blackish, often distinct. Inflorescences 14–57 mm long; glomerules distinct, at least at base distinctly interrupted, 1.5–2.5 by 1.5–2 mm |
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29b. |
Leaf: apex cuspidate to caudate; texture (rather) papery; dots greyish, often indistinct or fading. Inflorescences 5–23 mm long; glomerules distinct or not, distinctly or only slightly interrupted at base, 0.5–1.5 (seldom up to 2.5) by 0.3–1.5 mm, or indistinct and flowers seemingly continuously laxly set along the rachis |
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30a. |
Stipules falcate, early caducous or still present in young stages. Leaf: dots irregularly set, fine; midrib sparsely tomentose above. Inflorescences: glomerules distinctly interrupted throughout. Stamens 3 or 4 |
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30b. |
Stipules narrowly ovate, (slightly) oblique, early caducous. Leaf: dots densely set, large; glabrous above. Inflorescences: glomerules at base strongly interrupted, less at apex. Stamens 2 or 3 |
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31a. |
Stipules slightly asymmetric, mostly early caducous, sometimes present in young stages. Branchlets, petiole, and leaf only on midrib beneath sparsely puberulous |
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31b. |
Stipules slightly to strongly oblique, present in young stages or longer. Branchlets, petiole, and leaf at least on midrib above and beneath (sparsely) tomentose |
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32a. |
Petiole 7–15 mm long. Leaves drying rather shiny greenish on both sides, not blackening with age; nerves 7–10 pairs |
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32b. |
Petiole 4–8 mm long. Leaves drying above greenish to brown, beneath brown, blackening with age; nerves 4–6 pairs |
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33a. |
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33b. |
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34a. |
Leaf: base often slightly attenuate; drying greenish grey to greenish brown above, greyish brown beneath; nerves 6–8 pairs; nervation laxly reticulate, slightly scalariform. Inflorescences 5–9 mm long; glomerules c. 1 by 0.5 mm |
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34b. |
Leaf: base cuneate to emarginate; drying (dark) greenish brown above, brown beneath; nerves 8–17 pairs; nervation distinct, often distinctly reticulate-scalariform. Inflorescences 9–23 mm long; glomerules 1–2.5 by 1–1.5 mm |
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35a. |
Branchlets (sparsely) tomentose. Petiole 5–38 mm long, (sparsely) tomentose. Leaves at least (sparsely) tomentose beneath |
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35b. |
Branchlets sparsely puberulous. Petiole 14–51 mm long, sparsely puberulous. Leaves only on midrib sparsely puberulous beneath |
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36a. |
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36b. |
Stipules falcate. Petiole 8–38 mm long. Leaves: texture rather leathery; dots irregularly set, fine. Glomerules mostly distinctly interrupted throughout, globose, 1.5–2.5 by 1.5–2.5 mm |
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37a. |
Leaf: nerves 10–12 pairs; nervation laxly reticulate. Inflorescences 18–47 mm long; glomerules ellipsoid |
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37b. |
Leaf: nerves 8–17 pairs; nervation densely reticulate, (slightly) scalariform. Inflorescences 9–23 mm long; glomerules (sub)globose |
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1a. |
Ovary (mostly) 2-locular |
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1b. |
Ovary (mostly) 3- or more locular |
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2a. |
Septae and column (sparsely) hairy |
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2b. |
Septae and column glabrous |
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3a. |
Petiole and leaves at least sparsely puberulous beneath. Ovary (sparsely) tomentose. Fruit ovoid to ellipsoid, (sparsely) tomentose |
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3b. |
Petiole and leaves completely glabrous. Ovary glabrous. Fruit globose, glabrous |
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4a. |
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4b. |
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46c. A. octandra var. malesiana |
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5a. |
Ovary and fruit (sub)sessile, pedicel up to 0.5 mm long |
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5b. |
Ovary and fruit distinctly pedicelled, pedicel more than 1 mm long |
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6a. |
Stipules early caducous or present in young stages only, (narrowly) ovate, sometimes oblique. Ovary ellipsoid, densely to sparsely sericeous or puberulous; stigmas 0.5–1.5 mm long. Fruits narrowly ovoid or ellipsoid, seldom globose, not beaked, (sparsely) tomentose to subglabrous |
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6b. |
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7a. |
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7b. |
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8a. |
Leaves 11–25 by 5–11 cm. Inflorescences 10–42 mm long; stigmas flattened on top and to the sides of the ovary, densely laciniate. Infructescences 16–57 mm long |
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8b. |
Leaves 5.5–13 by 2–4.5 cm. Inflorescences 10–23 mm long; stigmas at first ascending from top to an apical bunch before flattening to the sides of the ovary, ribbed and papillate. Infructescences 12–33 mm long |
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9a. |
Leaf: nerves 6–10 pairs. Inflorescences 0.5–0.8 mm diam., (sparsely) sericeous. Ovary sparsely sericeous at base, glabrescent; stigma lowly papillate to laciniate. Fruits sparsely sericeous at base only, glabrescent, granular |
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9b. |
Leaf: nerves 8–17 pairs. Inflorescences 1.5–2 mm diam., tomentose. Ovary densely sericeous; stigmas densely laciniate. Fruits (sparsely) sericeous, punctate |
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10a. |
Stipules slightly asymmetric, early caducous, sometimes still present in young stages. Leaves drying rather shiny greyish green on both sides; marginal glands distinct, black to grey; above glabrous, beneath only midrib and nerves puberulous |
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10b. |
Stipules slightly to strongly oblique, present in young stages. Leaves drying greenish grey to greenish brown above, greyish brown beneath; marginal glands indistinct; above at least midrib sparsely puberulous, beneath sparsely puberulous |
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11a. |
Branchlets, petiole, and leaf completely glabrous. Leaves drying yellowish to greyish green on both sides. Ovary glabrous; stigmas 0.2–1 mm long. Fruits glabrous, drying yellowish; pericarp 0.2–1 mm thick |
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11b. |
Branchlets, petiole, and leaf at least on midrib beneath sparsely puberulous. Leaves drying dark greyish, greenish, bluish, or brownish above, brownish beneath. Ovary sericeous to sparsely puberulous at base only, if glabrous than stigmas longer than 1.5 mm; stigmas 0.8–5.5 mm long. Fruits sericeous to glabrous, drying dark to orange-brown; pericarp 0.5–1.5 mm thick |
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12a. |
Leaf: nerves 7–11 pairs, regularly set; dots indistinct. Flowers up to 11; ovary not lobed. Fruits ellipsoid or fusiform, 9–13 by 8–11 mm or 14–17 by 7–10 mm, drying smooth to punctulate; apex of endocarp acute |
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12b. |
Leaf: nerves 5–9 pairs, with often larger gap between 3rd and 4th pair; dots densely set, fine, greyish. Flowers up to 5; ovary lowly lobed. Fruits ovoid, 12–15 by 10–13 mm, often drying irregularly ridged and corrugated; apex of endocarp caudate |
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13a. |
Fruits fusiform, 14–17 mm long; pedicel 11–35 mm long |
66b. A. symplocoides var. chalarocarpa |
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13b. |
Fruits ovoid, 9–13 mm long; pedicel 2–7 mm long |
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14a. |
Leaf: dots indistinct or fading, if distinct then laxly irregularly set. Flower: pedicel 1–3 mm long; ovary (narrowly) ovoid; stigmas more or less raised, 1.5–5.5 mm long, free, perpendicular. Fruit (narrowly) ovoid to slightly flask-shaped or fusiform, slightly stiped and/or beaked |
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14b. |
Leaf: dots densely set, large, black. Flower: pedicel 0.8–1 mm long; ovary ellipsoid; stigmas sessile, 0.8–1.2 mm long. Fruit ellipsoid to subglobose, not stiped nor beaked |
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15a. |
Leaf: apex acuminate; drying greenish or reddish brown above; nerves 7–12 pairs. Ovary 2- and 3-locular; stigmas 1.5–2.5 mm long. Fruits not or only very slightly stiped, 13–22 mm long; pericarp 0.8–1.5 mm thick |
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15b. |
Leaf: apex cuspidate; drying dark greenish blue above; nerves 4–6 pairs. Ovary 2-locular, never 3-locular; stigmas 2.5–5.5 mm long. Fruits shortly stiped, 7–10 mm long; pericarp 0.5–0.8 mm thick |
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16a. |
Branchlets, petioles, and leaves only on midrib and nerves beneath very sparsely puberulous, glabrescent. Leaf: texture leathery; dots irregularly set, distinct. Ovary rugose, (sub)glabrous. Fruits (sub)glabrous; apex of endocarp acute |
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16b. |
Branchlets, petioles and leaves beneath (sparely) tomentose. Leaf: texture papery; dots often indistinct. Ovary smooth, (sparsely) sericeous. Fruits sparsely puberulous; apex of endocarp caudate |
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17a. |
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17b. |
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37d. A. lucida var. trilocularis |
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18a. |
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18b. |
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19a. |
Leaves glabrous or only sparsely hairy beneath; nerves slightly prominent to flat above |
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19b. |
Leaves at least on midrib and beneath (sparsely) hirsute above; nerves more or less sunken above |
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20a. |
Stipules 6–20 by 11–37 mm, often long persistent. Petiole upper pulvinus swollen, 5–11 by 3–6 mm. Leaves 14–47 cm long; base cordate; nerves beneath raised, not fading towards the margin |
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20b. |
Stipules 4–8 by 11–18 mm, present in young stages. Petiole upper pulvinus distinct, but not extremely so, 2–6 by 1.5–2.5 mm. Leaves 8–25 cm long; base emarginate to rounded; nerves beneath prominent, fading towards the margin |
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21a. |
Nerves and venation strongly impressed above and giving the leaf a bullate appearance |
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21b. |
Only nerves impressed, sometimes near margin and apex some veins impressed, leaves flat, smooth above |
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22a. |
Leaves 9–20 cm long |
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22b. |
Leaves 21–29 cm long |
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23a. |
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23b. |
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24a. |
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24b. |
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25a. |
Branchlets, petiole, and leaves on midrib and nerves beneath sparsely hairy. Inflorescences and flowers (sparsely) puberulous to tomentose. Infructescences and pedicels sparsely puberulous. Fruits (broadly) ovoid to globose, 8–12 mm long; apex of endocarp acute |
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25b. |
Branchlets, petiole, and leaves completely glabrous. Inflorescences and flowers subglabrous. Infructescences and pedicels glabrous. Fruits ovoid to ellipsoid, 13–21 mm long; apex of endocarp caudate |
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26a. |
Leaf: 5.5–13.5 cm long; dots irregularly densely set, greyish, fading. Stigmas slightly raised, narrow, not prominent. Fruits glabrous, drying brownish with light yellow-brown sutures; stigmas narrow, not extremely thick. — North Borneo, Mt Kinabalu and environments, above 1000 m |
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26b. |
Leaf: 10–19 cm long; dots absent. Stigmas sessile, broad, prominent. Fruits sometimes sparsely puberulous near base, drying light yellowish; stigmas broad, thick. — Sarawak, along or partly in rivers or tidal zones |
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27a. |
Branchlets and leaves (sparsely) hairy to glabrous. Leaf: nerves 8–13 pairs. Inflorescences sparsely tomentose to subglabrous. Ovary sparsely puberulous to glabrous. Fruits sparsely puberulous to glabrous |
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27b. |
Branchlets and leaves tomentose. Leaf: nerves 10–14 pairs. Inflorescences tomentose. Ovary puberulous. Fruits (sparsely) puberulous |
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28a. |
Branchlets and leaves (very) sparsely puberulous, glabrescent. Leaves 6.5–21 cm long, papery; nerves 8–11 pairs. Ovary mostly 3-locular, occasionally 4-locular, sparsely puberulous to glabrous. Fruits 9–18 mm long; pericarp 0.5–3(–6) mm thick |
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28b. |
Branchlets and leaves completely glabrous. Leaves 15–33 cm long, leathery; nerves 9–13 pairs. Ovary 4–6-locular, seldom 7-locular, glabrous. Fruits 21–28 mm long; pericarp 3–5 mm thick |
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29a. |
Branchlets and leaves beneath very sparsely puberulous, glabrescent. Inflorescences subglabrous. Ovary glabrous; stigmas papillate to laciniate. Infructescences subglabrous. Fruits young ones ridged, mature ones smooth, glabrous, stigmas not sunken |
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29b. |
Branchlets and leaves sparsely puberulous, glabrescent. Inflorescences (sparsely) tomentose. Ovary sparsely hairy, glabrescent; stigmas ribbed. Infructescences (sparsely) tomentose. Fruits young and mature ones smooth, sparsely hairy, glabrescent, stigmas flat to sunken Mostly A. frutescens is distinct by its stronger indument and smaller fruits from the typical subglabrous and large-fruited A. banahaensis. However, in places were A. banahaensis and A. frutescens co-occur, as in Sabah, the differences between the species fade: A. frutescens becomes glabrescent and the fruit of A. banahaensis does not get as large as in, e.g., Mindanao. |
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30a. |
Branchlets, petioles, and leaves completely glabrous. Leaves drying greyish green to yellowish green on both sides. Ovary glabrous; stigma up to 1 mm long. Fruits glabrous, yellowish when dry |
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30b. |
Branchlets, petioles, and leaves at least on midrib beneath very sparsely puberulous to tomentose. Leaves drying (dark) greyish green, bluish green, or brownish above, brownish beneath. Ovary at least at base sparsely puberulous (if (sub)glabrous, then stigmas 1.5–2.5 mm long, or nerves 14–23 pairs); stigmas 0.7–2.5 mm long. Fruits sparsely puberulous to glabrous, brownish to black when dry |
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31a. |
Leaves 6.5–18 cm long, smooth or young ones irregularly lowly granular beneath |
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31b. |
Leaves 13–32 cm long, strongly white-granular beneath on nervation only |
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32a. |
Leaf: nerves 7–11 pairs, regularly set; dots indistinct. Flowers up to 11; ovary not lobed. Fruits ellipsoid or fusiform, 9–13 by 8–11 mm or 14–17 by 7–10 mm, drying smooth to punctulate; apex of endocarp acute |
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32b. |
Leaf: nerves 5–9 pairs, with often larger gap between 3rd and 4th pair; dots densely set, fine, greyish. Flowers up to 5; ovary lowly lobed. Fruits ovoid, 12–15 by 10–13 mm, often drying irregular ridged and corrugate; apex of endocarp caudate |
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33a. |
Fruits fusiform, 14–17 mm long; pedicel 11–35 mm long |
66b. A. symplocoides var. chalarocarpa |
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33b. |
Fruits ovoid, 9–13 mm long; pedicel 2–7 mm long |
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34a. |
Stipules (narrowly) ovate, symmetric or slightly oblique, early caducous (if falcate and persistent, then nerves 8–13 pairs). Leaf: base rounded to cuneate, very base never cordate; nerves 6–16 pairs. Inflorescences sparsely puberulous to (sparsely) tomentose. Ovary (sparsely) puberulous to subglabrous. Fruits sparsely puberulous to glabrous; pericarp 0.8–5 mm thick |
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34b. |
Stipules falcate, often persistent. Leaf: base emarginate or cuneate with very base cordate; nerves 14–23 pairs. Inflorescences subglabrous. Ovary glabrous. Fruits glabrous; pericarp 0.5–1 mm thick |
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35a. |
Petiole 6–38 mm long. Leaf: texture papery or (slightly) leathery; disc-like absent, or few present along the margin; nerves 6–13 pairs. Inflorescences axillary to just below the leaves |
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35b. |
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36a. |
Branchlets, petioles, and leaves at least beneath (sparsely) tomentose to (sparsely) puberulous. Ovary smooth, at base (sparsely) tomentose to puberulous, glabrescent towards apex. Fruits sparsely puberulous |
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36b. |
Branchlets, petioles, and leaves only on midrib and nerves beneath very sparsely puberulous, glabrescent. Ovary rugose, (sub)glabrous. Fruits (sub)glabrous |
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37a. |
Leaf: dots often distinct, large or small. Ovary ovoid or ellipsoid; stigmas sessile, 0.8–2.2 mm long. Fruits ellipsoid to (sub)globose, not stiped or beaked; apex of endocarp acute |
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37b. |
Leaf: dots indistinct. Ovary narrowly ovoid; stigmas slightly raised, 2–2.5 mm long. Fruits ovoid to slightly flask-shaped, beaked, 13–19 mm long; apex of endocarp caudate |
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38a. |
Leaf: dots irregularly set, fine; nerves 8–13 pairs. Stipules falcate, sometimes persistent. Flowers: sepals 3; ovary 3-locular (seldom 4-locular); stigmas 1.5–2.2 mm long. Fruits: pericarp 1–5 mm thick |
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38b. |
Leaf: dots laxly set, large; nerves 6–9 pairs. Stipules caducous. Flowers: pedicel 0.8–1 mm long; ovary 2- and 3-locular; stigmas 0.8–1.2 mm long. Fruits: pericarp 0.8–1.5 mm thick |