Key to the Species of Borneo

 

Key to Pistillate specimens

 

Staminate specimens

 

1a.

Staminate inflorescences strongly interrupted; all individual glomerules distinct, widely set along the rachis (click picture)

2

1b.

Staminate inflorescences continuous to more or less interrupted at base; individual glomerules at least at apex of rachis indistinct, not or only slightly spaced (click picture)

11

2a.

Branchlets, petioles and leaves completely glabrous

3

2b.

Branchlet, petioles and leaves on midrib beneath sparsely puberulous

6

3a.

Leaves 6.5–18 cm long, smooth or young ones irregularly lowly granular beneath

4

3b.

Leaves 13–32 cm long, strongly white-granular beneath on nervation only

14. A. chondroneura

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

5

4b.

Leaf: nerves 9–12 pairs, drying often beneath lighter green than the lamina. Inflorescences: glomerules apically less spaced than at base

56. A. sarawakensis

5a.

Leaf: nerves 5–8 pairs; texture slightly leathery, drying dull greyish green to dark green; dots densely set, greyish

51. A. prainiana

5b.

Leaf: nerves 7–10 pairs; texture papery, drying shiny yellowish green or dark green; dots indistinct

66. A. symplocoides

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

7

6b.

Stipules falcate, persistent. Leaf: base emarginate-truncate; nerves 14–23 pairs

25. A. grandistipula

7a.

Branchlets sparsely tomentose to subglabrous. Leaves only sparsely tomentose to subglabrous beneath; drying yellowish green on both sides

8

7b.

Branchlets tomentose. Leaf: at least (sparsely) tomentose beneath; drying dark brownish to greenish brownish above, brown beneath

9

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

23. A. fulvovittata

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

62. A. stenostachys

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

10

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

29. A. illustris

10a.

Leaves 11–25 by 5–11 cm. Inflorescences 10–20 clustered together, 11–47 mm long

19. A. elmeri

10b.

Leaves 5.5–13 by 2–4.5 cm. Inflorescences 2–5 clustered together, 7–12 mm long

55. A. rhacostyla

11a.

Staminate rachis continuously floriferous from apex to base, though individual glomerules sometimes minutely spaced; peduncle absent, seldom minute (click picture)

12

11b.

Staminate rachis at least at base free of flowers; peduncle more than 1 mm long (click picture)

14

12a.

Petiole and leaf at least beneath puberulous. Inflorescences continuous, glomerules uninterrupted, indistinct

13

12b.

Petiole and leaf completely glabrous. Inflorescences continuous, glomerules slightly interrupted, distinct

37. A. lucida

13a.

Leaves 12–28 cm long; margin (lowly) glandular-crenate (click picture); only sparsely hirsute beneath; drying green or brown, blackening with age; disc-like glands absent

43. A. nigricans

13b.

Leaves 5–16 cm long; margin more or less distinctly glandular-serrate (click left picture); (sparsely) puberulous on both sides; drying brownish or yellowish green, not blackening with age; disc-like glands many, scattered in centre of marginal vein arches (click right picture)

46c. A. octandra var. malesiana

14a.

Stipules falcate, persistent or caducous. Leaf: marginal glands in margin, small and indistinct; disc-like glands often present, distinct, along the margin (click upper left picture); nerves 9–18 pairs. Inflorescences (when mature) brush-like due to exserted stamens, glomerules indistinct, densely set at apical 3/4–7/8 of the rachis (click lower picture). Flowers: stamens strongly exserted, c. thrice as long as the sepals (click upper right picture); anthers 0.1–0.2 mm long

15

14b.

Stipules narrowly ovate, oblique or not, seldom falcate (but then leaf nerves up to 9 pairs or inflorescences interrupted), caducous but sometimes still present in young stages. Leaf: marginal glands often rather distinct; disc-like glands absent or occasionally present, then indistinct, along margin (click upper left picture) , seldom scattered at base within marginal vein arches (click upper right picture). Inflorescences (when mature) not brush-like, glomerules at least at base distinct, slightly to strongly interrupted (if glomerules continuous and indistinct, then either flowers laxly set or stamens not exserted (click lower left picture) or inflorescences clavate-shaped) (click lower richt picture). Stamens not to slightly exserted (up to twice as long as the sepals), if exserted, then anthers 0.3–0.6 mm long

19

15a.

Leaves glabrous or only beneath sparsely hairy; nerves slightly prominent to flat above

16

15b.

Leaves: above at least on midrib (sparsely) hirsute, beneath (sparsely) hirsute; nerves more or less sunken above

17

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

8. A. benthamiana

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

20. A. falcifera

17a.

Nervation strongly sunken above, giving the leaf a bullate appearance

18

17b.

Only nerves sunken, sometimes near margin and apex some veins sunken, otherwise leaves flat, smooth above

38. A. lunata

18a.

Leaves 9–20 cm long

11. A. bullatissima

18b.

Leaves 21–29 cm long

64. A. sylvestri

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

20

19b.

Branchlets and petiole (sparsely) puberulous; leaves at least on midrib beneath (sparsely) puberulous to tomentose. Inflorescences: rachis (sparsely) puberulous to tomentose

25

20a.

Leaves 6–20 by 1.5–8 cm. Inflorescences cylindric or interrupted (click picture), 5–41 mm long, 0.8–3.5 mm diameter

21

20b.

Leaves 15–33 by 5–13 cm. Inflorescences clavate (click picture), 12–36 mm long, apical head 3–8 mm diameter

45. A. nitida

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

22

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

3. A. antennifera

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

23

22b.

Leaf: texture leathery, shiny; drying greenish to brownish above, brownish beneath; dots often irregularly set, distinct, but often fading. Glomerules 2–3 mm long. Flowers drying brownish; stamens exserted, c. twice as long as sepals (click picture)

12. A. caloneura

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

24

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

56. A. sarawakensis

24a.

Inflorescences 1–1.5 mm diam.; glomerules rather distinctly interrupted at base of rachis (click picture)

6. A. banahaensis

24b.

Inflorescences 2–3.5 mm diam.; glomerules at base only slightly interrupted (click picture)

22. A. frutescens

25a.

Inflorescences 1–6 clustered together; glomerules subglobose to ellipsoid, distinct to indistinct, (slightly) interrupted at base, decreasingly spaced towards the apex

26

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

62. A. stenostachys

26a.

Leaves drying greyish green to yellowish green. Inflorescences 2–9 mm diam.; glomerules clustered strongly apically on fleshy rachis (click picture); drying yellowish

27

26b.

Leaves drying greenish or brownish. Inflorescences 0.5–2.2 mm diam.; glomerules not clustered densely apically (click picture); drying light to dark brown

28

27a.

Branchlets and leaves tomentose. Leaf: nerves 10–14 pairs. Inflorescences 12–32 mm long, tomentose; apical head 5–9 mm diameter

15. A. confusa

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

22. A. frutescens

28a.

Leaves: nerves 4–10 pairs

29

28b.

Leaves: nerves 10–17 pairs

35

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

30

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

31

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 

29. A. illustris

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

42. A. nervosa

31a.

Stipules slightly asymmetric, mostly early caducous, sometimes present in young stages. Branchlets, petiole, and leaf only on midrib beneath sparsely puberulous

32

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

33

32a.

Petiole 7–15 mm long. Leaves drying rather shiny greenish on both sides, not blackening with age; nerves 7–10 pairs

1. A. alia

32b.

Petiole 4–8 mm long. Leaves drying above greenish to brown, beneath brown, blackening with age; nerves 4–6 pairs

3. A. antennifera

33a.

Stipules slightly to strongly oblique, caducous, present in young stages only. Leaves above only on midrib (sparsely) puberulous to (sparsely) tomentose. Inflorescences: glomerules at least at base distinct, globose, compact (click picture); stamens not to slightly exserted

34

33b.

Stipules strongly oblique, often rather long persistent. Leaves above sparsely hirsute. Inflorescences: glomerules lax, indistinct, flowers seemingly continuously laxly set along the rachis (click picture); stamens exserted

30. A. lagenocarpa

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

26. A. granularis

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

63. A. subcaudata

35a.

Branchlets (sparsely) tomentose. Petiole 5–38 mm long, (sparsely) tomentose. Leaves at least (sparsely) tomentose beneath

36

35b.

Branchlets sparsely puberulous. Petiole 14–51 mm long, sparsely puberulous. Leaves only on midrib sparsely puberulous beneath

4. A. arborea

36a.

Stipules narrowly ovate, sometimes oblique. Petiole 5–17 mm long. Leaves: texture (rather) papery; dots indistinct or fading. Glomerules interrupted at base only, rather continuously set apically (click picture), subglobose or ellipsoid, 1–2.5 by 1–1.5 mm

37

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

29. A. illustris

37a.

Leaf: nerves 10–12 pairs; nervation laxly reticulate. Inflorescences 18–47 mm long; glomerules ellipsoid

7. A. basilanensis

37b.

Leaf: nerves 8–17 pairs; nervation densely reticulate, (slightly) scalariform. Inflorescences 9–23 mm long; glomerules (sub)globose

63. A. subcaudata

 

Pistillate specimens

 

1a.

Ovary (mostly) 2-locular

2

1b.

Ovary (mostly) 3- or more locular 

17

2a.

Septae and column (sparsely) hairy

3

2b.

Septae and column glabrous

5

3a.

Petiole and leaves at least sparsely puberulous beneath. Ovary (sparsely) tomentose. Fruit ovoid to ellipsoid, (sparsely) tomentose

4

3b.

Petiole and leaves completely glabrous. Ovary glabrous. Fruit globose, glabrous

37a. A. lucida var. lucida

4a.

Leaves 12–28 cm long; margin (lowly) glandular-crenate (click picture); only sparsely hirsute beneath; drying green or brown, blackening with age; disc-like glands absent. Fruits drying dark brown to black with light sutures

43. A. nigricans

4b.

Leaves 5–16 cm long; margin more or less distinctly glandular-serrate (click left picture); (sparsely) puberulous on both sides; drying brownish or yellowish green, not blackening with age; disc-like glands many, scattered within marginal vein arches (click right picture). Fruits drying orange-brown

46c. A. octandra var. malesiana

5a.

Ovary and fruit (sub)sessile, pedicel up to 0.5 mm long 

6

5b.

Ovary and fruit distinctly pedicelled, pedicel more than 1 mm long 

11

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

7

6b.

Stipules often rather persistent, narrowly ovate, strongly oblique to falcate. Ovary narrowly ovoid, flask-shaped to subfusiform, densely tomentose; stigmas 4.5–6 mm long (click left picture). Fruits flask-shaped to subfusiform, beaked, (sparsely) tomentose (click right picture)

30. A. lagenocarpa

7a.

Stipules (narrowly) ovate, symmetric. Flowers and fruits rather laxly set along the whole rachis (click picture)

8

7b.

Stipules narrowly ovate, more or less oblique. Flowers and fruits rather densely set at least at apical part of the rachis (click picture)

9

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

19. A. elmeri

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 

55. A. rhacostyla

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 

10

9b.

Leaf: nerves 8–17 pairs. Inflorescences 1.5–2 mm diam., tomentose. Ovary densely sericeous; stigmas densely laciniate. Fruits (sparsely) sericeous, punctate

63. A. subcaudata

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

1. A. alia

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

26. A. granularis

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

12

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

14

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

13 (66. A. symplocoides)

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

52. A. prainiana

13a.

Fruits fusiform, 14–17 mm long; pedicel 11–35 mm long

66b. A. symplocoides var. chalarocarpa

13b.

Fruits ovoid, 9–13 mm long; pedicel 2–7 mm long

66a. A. symplocoides var. symplocoides

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

15

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

42. A. nervosa

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

16

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

3. A. antennifera

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

12. A. caloneura

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

7. A. basilanensis

17a.

Leaf: disc-like glands absent or only at base (click upper left picture) or along the margin (click upper right pictures). Flowers and fruits densely set only at apical part of rachis (click lower left picture), or laxly set along the rachis (click lower right picture) ; subsessile to distinctly pedicelled. Fruit: septae and column glabrous

18

17b.

Leaf: disc-like glands scattered within the marginal vein arches (click left picture). Flowers and fruits densely set along the rachis (click right pictures), sessile. Fruit: septae and column sparsely sericeous

37d. A. lucida var. trilocularis

18a.

Flowers and fruits subsessile, densely set only along apical 7/8–3/4 of the rachis (click left picture). Stigmas sessile, smooth, flattened to the sides of the ovary (click central picture). Fruits ovoid with stigma remnant ascending. Leaf: disc-like glands distinct, along the margin (click right picture)

19

18b.

Flowers and fruits distinctly pedicelled, set rather laxly along the whole rachis (click upper left picture) or singly apically (click upper right picture). Stigmas sessile to raised, ribbed to laciniate, perpendicular to, or ascending from the sides of the ovary. Fruits ellipsoid to globose, if ovoid, then distinctly pedicelled and stigma remnants usually not ascending. Leaf: disc-like glands absent, or only a few distinct ones along the margin, sometimes few scattered near base within the marginal vein arches (click lower pictures)

23

19a.

Leaves glabrous or only sparsely hairy beneath; nerves slightly prominent to flat above

20

19b.

Leaves at least on midrib and beneath (sparsely) hirsute above; nerves more or less sunken above

21

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

8. A. benthamiana

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

20. A. falcifera

21a.

Nerves and venation strongly impressed above and giving the leaf a bullate appearance

22

21b.

Only nerves impressed, sometimes near margin and apex some veins impressed, leaves flat, smooth above

38. A. lunata

22a.

Leaves 9–20 cm long

11. A. bullatissima

22b.

Leaves 21–29 cm long

64. A. sylvestri

23a.

Inflorescences and infructescences bearing usually only one single flower or fruit respectively on top of the rachis (click picture) (seldom few clustered apically). Stigmas broad, raised or sunken (if narrow, then altitude > 1000 m, Mt Kinabalu or other mountains in northern Borneo). — Fruit sometimes globose with sunken stigmas and thick fleshy pericarp

24

23b.

Inflorescences and infructescences bearing more than one flower or fruit respectively along the rachis (click picture). Stigmas narrow

30

24a.

Fruits ellipsoid or (broadly) ovoid with prominent stigmas, seldom subglobose; pericarp 0.5–1.5 mm thick. Ovary ovoid, stigmas slightly raised, ribbed and margins laciniate, or cylindric and stigmas broad and prominent (click picture); stigmas 1–2 mm long

25

24b.

Fruits globose with sunken stigmas (click left picture), seldom ovoid, but then sutures ridged; pericarp 0.5–6 mm thick. Ovary globose, stigmas broad, flat, 0.5–1 mm long, slightly ribbed or laxly laciniate (click right picture)

27

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

26

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

56. A. sarawakensis

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

23. A. fulvovittata

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

62. A. stenostachys

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

28

27b.

Branchlets and leaves tomentose. Leaf: nerves 10–14 pairs. Inflorescences tomentose. Ovary puberulous. Fruits (sparsely) puberulous

15. A. confusa

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

29

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

45. A. nitida

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

6. A. banahaensis

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.

22. A. frutescens

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

31

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

34

31a.

Leaves 6.5–18 cm long, smooth or young ones irregularly lowly granular beneath

32

31b.

Leaves 13–32 cm long, strongly white-granular beneath on nervation only

14. A. chondroneura

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

33 (66. A. symplocoides)

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

51. A. prainiana

33a.

Fruits fusiform, 14–17 mm long; pedicel 11–35 mm long

66b. A. symplocoides var. chalarocarpa

33b.

Fruits ovoid, 9–13 mm long; pedicel 2–7 mm long

66a. A. symplocoides var. symplocoides

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

35

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

25. A. grandistipula

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

36

35b.

Petiole 14–51 mm long. Leaf: texture papery; disc-like glands often present, along the margin (click picture); nerves 12–16 pairs. Inflorescences below the leaves or on the branches

4. A. arborea

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 

37

36b.

Branchlets, petioles, and leaves only on midrib and nerves beneath very sparsely puberulous, glabrescent. Ovary rugose, (sub)glabrous. Fruits (sub)glabrous

12. A. caloneura

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

38

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

7. A. basilanensis

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

29. A. illustris

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

42. A. nervosa