Antipodanthus, Bouman & Keβler & Telford & Bruhl & Welzen, 2018

Bouman, R. W., Keβler, P. J. A., Telford, I. R. H., Bruhl, J. J. & Welzen, P. C. van, 2018, Subgeneric delimitation of the plant genus Phyllanthus (Phyllanthaceae), Blumea 63 (2), pp. 167-198 : 170-175

KEY TO THE SUBGENERA AND (SUB)SECTIONS OF PHYLLANTHUS

A provisional key is here provided based on characters mentioned in the literature. A key for full identification purposes, using morphology only (not pollen) is difficult due to the absence of recent complete treatments for several groups and the fact that some characters have evolved multiple times within Phyllanthus . The key is not completely dichotomous (trichotomous questions are marked with *). Authors of the various subgenera, sections and subsections are listed in Appendix 1 and all species within a particular group are listed in Appendix 2.

1. Branching non-phyllanthoid (laminate leaves and flowers on all axes, branchlets not deciduous)................. 2

1. Branching phyllanthoid (leaves on main stem reduced to scales, cataphylls, laminate leaves and flowers on lateral axes, lateral branchlets deciduous) or sub-phyllanthoid (leaves at base of branchlets not reduced to scales (often in juveniles), lateral branchlets deciduous).......... 15

2. Aquatic herbs. — Americas.................................... Subgenus Phyllanthus section Salviniopsis

2. Herbs, shrubs or trees, but not aquatic.............. 3

3. Palm-like (monocaul) shrubs to trees; stigma petaloid. — South America.. Subgenus Xylophylla section Asterandra

3. Herbs, shrubs to small trees, rarely climbers; stigmas vari- ously bifid to multifid, not petaloid.................. 4

4. Leaves on all axes spirally arranged................ 5

4. Leaves on all axes distichous..................... 9

5. Sepals 4 in staminate flowers, 6 in pistillate flowers; staminate disc entire, H-shaped around filaments; stamens 2, filaments free. — North America................................ Subgenus Swartziani section Reverchonia

5. Sepals 5 or 6 in both sexes; staminate disc segmented; stamens 3 or 5, filaments free or connate............ 6

6. Sepals 5; stamens 5, filaments free. — Africa....................Subgenus Kirganelia section Pseudomenarda

6. Sepals 5 or 6; stamens usually 3, filaments connate (free in P. rosmarinifolius Müll.Arg. )....................... 7

7. Inflorescences axillary cymules with 1–4 flowers; pistillate disc consisting of free glands. — Africa and Asia, introduced in North America................ Subgenus Isocladus

7. Inflorescences axillary cymules or thyrses; pistillate disc entire....................................... 8

8. Inflorescences axillary glomerules; pollen 3- or 4-colporate, subglobose. — South America and Australia?.................. Section Antipodanthus incertae sedis

8. Inflorescences axillary glomerules or thyrses (sometimes paniculate at end of branch); pollen areolate. — Central and South America...... Subgenus Xylophylla section.................................... Elutanthos

9. Staminate disc segmented, pistillate disc entire or segmented; filaments free........................ 10

9. Staminate disc segmented, or entire and urceolate, pistillate disc often massive and urceolate; filaments connate. Pollen with macroreticulate exine....................................... 12 (Subgenus Ceramanthus )

10. Anthers dehiscing with horizontal slits; pollen 4-colporate. — Americas........................................ Subgenus Phyllanthus section Loxopodium

10. Anthers often deflexed, but dehiscing with vertical slits; pollen clypeate or perisyncolporate.............. 11

11. Leaves distichous; pollen clypeate; seeds verrucate. — Africa, Asia, Australia and Pacific.. Subgenus Macraea

11. Leaves spiral at basal nodes, distichous at upper nodes; pollen grains perisyncolporate with median pores, colpi bordered by parallel muri; seeds smooth or striate. — Madagascar.................. Subgenus Betsileani

12. Staminate flowers with sepals 4 (6 in pistillate ones); staminate disc consisting of 4 massive segments; stamens 2, filaments connate and thecae on an enlarged connective. Pollen stephanoporate. — Africa................................ Subgenus Ceramanthus section Bivia

12. Sepals 6 in both sexes; staminate disc entire or 6 segments; stamens 3 with connate filaments, thecae not on an enlarged connective....................... 13

13. Staminate disc entire and cup-shaped. — Asia.............. Subgenus Ceramanthus section Ceramanthus

13. Staminate disc segmented or only slightly fused into a ring....................................... 14

14. Sepals in two dimorphic whorls; staminate disc segmented; pollen peribrevicolporate. — Africa and Asia................. Subgenus Ceramanthus section Anisolobium

14. Sepals in two equal whorls; staminate disc segmented to slightly fused into a ring; pollen pantoporate. — Asia............. Subgenus Ceramanthus section Cluytopsis

15. Branching sub-phyllanthoid.................... 16

15. Branching phyllanthoid (sometimes sub-phyllanthoid in very young plants, check mature plants).......... 20

16. Branchlets short, with only 5–10 leaves ( Webster 2001b); staminate disc entire. — South and Central America............. Subgenus Xylophylla section Brachycladus

16. Branchlet length variable, usually bearing more than 10 leaves; staminate disc segmented.............. 17

17. Anther connective not enlarged; fruit an indehiscent capsule; seeds smooth with fleshy sarcotesta. — South America and West Indies...................................... Subgenus Conami section Hylaeanthus

17. Anther connective variable, often enlarged; fruit a dehiscent capsule; seeds ornamented, without a fleshy sarcotesta...................................... 18

18. Filaments connate, stamens mostly 3................................... 26 (subgenus Afroswartziani )

18. Filaments free (filaments connate in P. allemii G.L.Webster and P. fastigiatus Mart ex Müll.Arg. , but then only 2 stamens)..................................... 19

19. Anther connective often enlarged, thecae not appearing as stipitate; seeds scalariform with slight transverse striations or smooth. — Australia and Central America (?).................. Subgenus Phyllanthus section Lysiandra

19. Anther connective variable, sometimes deeply emarginate with the two thecae appearing to be stipitate; seeds striate or linearly verrucate. — South America....................... Subgenus Phyllanthus section Phyllanthus ......................... subsection Clausseniani

20. Herbs or subshrubs.......................... 21

20. Shrubs to trees, rarely climbers................. 36

21. Herbs; each branchlet bearing just one pair of (sub)opposite leaves and terminating in a raceme; anther connective enlarged. — South America................................Subgenus Phyllanthus section Almadanses

21. Herbs or subshrubs; branchlet with more than 2 alternate leaves and flowers in leaf axils; anther connective (not) enlarged................................... 22

22. Flowers 5-merous; stamens 5, filaments free (except 3 stamens in P. cocumbiensis Jean F.Brunel ). Pollen subglobose, 3- or 4-colporate. — Pantropical, but origin Africa...................... 23 (Subgenus Tenellanthus )

22. Flowers 5- or 6-merous; stamens 2 or 3, filaments free or connate.................................... 24

23*. Shrubs or hemicryptophytes; stamens 5, filaments basally united; pollen 3-colporate, with macroreticulate exine. Seeds with fine punctuation. — Africa......................... Subgenus Tenellanthus section Loandani

23*. Herbs; stamens 5 (3 in P. cocumbiensis ), filaments connate or free; pollen 3-colporate, with tectate, microperforate exine. — Africa......................................... Subgenus Tenellanthus section Pentandra

23*. Herbs or subshrubs; stamens 5, filaments completely free; pollen 4-colporate with sponge-like exine. — Pantropical, but origin Africa....................................... Subgenus Tenellanthus section Tenellanthus

24. Inflorescences unisexual...................... 25

24. Inflorescences bisexual....................... 35

25. Cataphyllary stipules (unilaterally) auriculate....... 26

25. Cataphyllary stipules not auriculate.............. 32

26. Leaf base symmetric; plagiotropic branches carinate (wing- ed) ( Brunel & Roux 1981). Pollen exine tectate; seeds with longitudinal striae or smooth. — Africa..................... Subgenus Afroswartziani section Odontadenii

26. Leaf base asymmetric; plagiotropic branches not carinate....................................... 27

27. Pistillate disc entire with delicate fringes. — Africa............... Subgenus Afroswartziani section Callidisci

27. Pistillate disc entire, but not fringed.............. 28

28. Sepals 5 in staminate flowers.................. 29

28. Sepals 6 in staminate flowers.................. 30

29. Cataphyllary stipules usually black and indurate; stamen 2 or 3, filaments partially or wholly connate, anthers sometimes deeply emarginate, dehiscing mostly horizontal; seeds longitudinally striate or banded, possibly with transverse striae. Pollen 3-colporate. — West Indies............. Subgenus Phyllanthus section Phyllanthus ......................... subsection Pentaphyllus

29. Cataphyllary stipules thin and membranous, not indurate or black; stamen 3, filaments connate, anthers not emarginate, dehiscing horizontally to vertically; seeds longitudinally striate.......................... 30

30. Pistillate inflorescences on proximal position and staminate inflorescences on distal position of plagiotropic branchlets; seeds transversely striate. Ovary often cov- ered with tubercles. — Pantropical, but origin Asia.................... Subgenus Emblica section Urinaria

30. Pistillate inflorescences on distal position and staminate inflorescences on proximal position of plagiotropic branchlets; seeds longitudinally striate....................................... 31 (Subgenus Afroswartziani )

31*. Pollen 3- or 4-colporate, exine bireticulate. — Africa........ Subgenus Afroswartziani section Praephyllanthus

31*. Pollen 3-sulcate, exine macroreticulate. Often found in water. — Africa...................................... Subgenus Afroswartziani section Fluitantoides

31*. Pollen perihexabrevisulcate, exine macro-rugulose ( Brunel 1987). Ovary on gynophore. — Africa.................... Subgenus Afroswartziani section Microdendron

32. Branchlets and flowers not purplish; stamens 3, filaments mostly free or united to 2/3 of length; pollen 4-colporate, exine (hetero-)reticulate; pistillate sepals 5; pistillate disc entire; stigmas free, bifid, tips sometimes subcapitate. 33

32. Branchlets and flowers often purplish; stamens 2 or 3, filaments connate; pollen pantoporate, exine shields elongated or if round with only 1 pila; pistillate sepals 6; pistillate disc dissected or lobed; stigmas free or connate, bifid to emarginate, tips not capitate 34 (Subgenus Cyclanthera )

33. Anther connective not enlarged, thecae not stipitate; style branches sub-capitate; seeds verrucate. — South America, pantropically invasive. Subgenus Phyllanthus ............... section Phyllanthus subsection Niruri

33. Anther connective variable, deeply emarginate with the two thecae appearing stipitate; style branches not capitate; seeds striate or linearly verrucate. — South America............. Subgenus Phyllanthus section Phyllanthus ......................... subsection Clausseniani

34. Branchlets unramified, rooting at nodes; leaves crisply succulent; stamens 2, filaments free; pollen shields elongated (banded) ( Webster & Carpenter 2002). — West Indies......... Subgenus Cyclanthera section Callitrichoides

34. Branchlets often with 1 or 2 lateral branches (bipinnatiform), not rooting at nodes; leaves not succulent; stamens 3, filaments completely connate into a circular synandrium; pollen shields isodiametric, each with a central pila surrounded by a murus ( Webster & Carpenter 2002). — West Indies..... Subgenus Cyclanthera section Cyclanthera

35. Stamens 3, filaments free, anthers dehiscing horizontally; pistillate disc dissected; pollen grains brevicolporate and diorate or porate, exine pilate; seeds verruculose. — South America........ Subgenus Conami section Apolepsis

35. Stamens 2 or 3, filaments entirely or partially connate (free in P. warnockii G.L.Webster ), anthers dehiscing oblique to horizontally (vertically in P. warnockii ); pistillate disc entire; pollen 3-colporate, exine reticulate; seeds smooth or longitudinally striate. — North America, pantropical invasive........................... Subgenus Swartziani

36. Leaves reduced and branchlets transformed to phylloclades (at least in older branches)............... 37

36. Leaves not reduced and branchlets not transformed to phylloclades................................ 39

37. All stems rounded or flat; stipules unilaterally auriculate, stamens free or connate; pollen 3- or 4-colporate, exine reticulate 38 (Subgenus Phyllanthus section Choretropsis )

37. Lateral stems flattened with wide phylloclades, (bi-)pinnatiform;stipules not auriculate;stamens usually united at base; pollen clypeate, exine areolate. — West Indies................... Subgenus Xylophylla section Xylophylla

38. Main axes often flat, branching monopodial, leaves distichous;inflorescences usually bisexual,stamens 3 (rarely 4). — South America............ Subgenus Phyllanthus .......... section Choretropsis subsection Applanata

38. Main axes rounded, branching monopodial or sympodial, leaves spiral; inflorescences mostly unisexual; stamens 2 or 3. — South America...... Subgenus Phyllanthus ........ section Choretropsi s subsection Choretropsis

39. Fruits indehiscent, berries or drupes............. 40

39. Fruits dehiscent, capsules (or absent)............ 48

40. Fruits drupaceous............................ 41

40. Fruits baccate............................... 43

41. Sepals 6; stamens 3, filaments connate. — Asia..................... Subgenus Emblica section Emblica

41. Sepals 4–6; stamens 3 or 4 (rarely 2 or 5), filaments free............ 42 (Subgenus Kirganelia section Cicca )

42. Plants dioecious; disc absent in both sexes; staminodes absent; fruits spongy ( Webster 1957). — West Indies and South America).... Subgenus Kirganelia section Cicca .......................... subsection Aporosella

42. Plants monoecious; disc present in both sexes; sometimes staminodes present; fruits hard. — Commonly cultivated, origin possibly African?..........Subgenus Kirganelia .............. section Cicca subsection Cheramella

43. Stamen 2, filaments connate; ovary 2-locular. — Africa, Madagascar, Mainland Asia................................. Subgenus Kirganelia section Chorisandra

43. Stamen 3‒6, filaments free or connate; ovary 3-locular .......................................... 44

44. Branchlets subtended by reduced leaves, but not cataphylls, flowers on brachyblasts; stamens 3, filaments free or connate. — South America................................. Subgenus Conami section Hylaeanthus

44. Branchlets subtended by (spinescent) cataphylls, stamens 4‒6, filaments free........................... 45

45. Branchlets subtended by spinescent cataphylls; stamens 5 in 2 sets, one free and the other basally fused. — Africa and Asia..... Subgenus Kirganelia section Anisonema

45. Branchlets subtended by scale or stipule like cataphylls; stamens 4–6, filaments free.................... 46

46. Staminate inflorescences on separate (leafless) plagiotropic branches, pistillate flowers axillary. Seeds globular, smooth. — Africa Subgenus Kirganelia section Polyanthi

46. Inflorescences axillary, on all plagiotropic branches.. 47

47. Stamens 5. — Asia..........................................Subgenus Kirganelia section Hemicicca

47. Stamens 6. — Africa, Madagascar, Mainland Asia................ Subgenus Kirganelia section Chorisandra

48. Anthers apiculate............................ 49

48. Anthers non-apiculate........................ 56

49. Sepals often caudate-acuminate; filaments connate, staminate disc consisting of linear spathulate segments; pistillate disc entire. Pollen 4-colporate, exine reticulate. — Asia............. 50 (Subgenus Phyllanthodendron )

49. Sepals often acuminate, but not caudate; filaments free or connate, staminate disc segmented, globular; pistillate disc entire (or absent)......................... 54

50. Shoots not differentiated, all leaves similar in size, flowers on lateral shoots............................. 51

50. Shoots differentiated into sterile leaf bearing shoots with larger leaves and fertile shoots with smaller leaves.. 53

51. Sepals 4 in staminate flowers; stamens 4. — Asia.....

..... Subgenus Phyllanthodendron section Tetrandrum 51. Sepals 5 or 6 in staminate flowers; stamens 3...... 52

52. Shrubs; sepals 5 in staminate flowers. Fruit reminiscent of Actephila ( Croizat 1942) . — Asia...................Subgenus Phyllanthodendron section Pseudoactephila

52. Twining shrubs; Sepals 6 sepals in staminate flowers. — Asia Subgenus Phyllanthodendron section Arachnodes

53. Trunk often succulent and enlarged at base; leaf blades> 6 cm long. — Asia............................ Subgenus Phyllanthodendron section Phyllanthodendron

53. Trunk not succulent or enlarged at base; leaf blades <6 cm long. — Asia..................................... Subgenus Phyllanthodendron section Calophyllum

54. Filaments connate. — South America..........................Subgenus Xylophylla section Ciccastrum

54. Filaments free............................... 55

55. Leaves with or without laminar glands; sepals in two indistinct whorls; pollen 4–8-colporate or diorate. — South America... Subgenus Emblica section Microglochidion

55. Leaves without laminar glands; sepals in two distinct whorls; pollen 3-syncolporate..... 67 (Subgenus Gomphidium )

56. Leaves opposite or subopposite................. 57

56. Leaves alternate............................. 59

57. Bark lenticellate; filaments connate. — Cuba............. Subgenus Xylophylla section Williamia subsection....................................... Mirifici

57. Bark smooth; filaments free.................... 58

58. Branchlets sometimes opposite bipinnatiform; staminate sepals 5, not distinctly biseriate; staminate disc consisting of 5 free segments; stamens 5. — Madagascar and Middle East (?)...................... Subgenus Menarda

58. Branchlets pinnatiform, not opposite; staminate sepals 5 or 6, in both sexes often distinctly biseriate; staminate disc entire, 3 emarginate segments or 5 or 6 massive segments; stamens mostly 3 or 5 (up to 20)............................... 67 (Subgenus Gomphidium )

59. Sepals 4 in staminate flowers; stamen 2, filaments connate. Pollen pantoporate or clypeate............. 60

59. Sepals 5 or 6 in staminate flowers; stamens 3–15, filaments free or connate......................... 66

60. Leaf margins very thick, conspicuously revolute; staminate disc massive, entire; pollen clypeate. — Cuba............... Subgenus Xylophylla section Glyptothamnus

60. Leaf margins not thickened, sometimes slightly revolute; staminate disc segmented; pollen pantoporate or clypeate....................................... 61

61. Anthers dehiscing vertically; sepal margins entire... 62

61. Anthers dehiscing horizontally/transversely; sepal margins entire to dentate or lacerate.................... 64

62. Inflorescences usually bisexual, appearing with the expanding leaves ( Webster 1958); pollen clypeate; style connate in a tube and stigmas often reduced to acute tips. — West Indies......... Subgenus Xylophylla section Thamnocharis

62. Inflorescences mostly unisexual, appearing after the leaves; pollen pantoporate; style connate or free... 63

63. Ovary papillose or verruculose, 3-locular. — Asia.............. Subgenus Eriococcus section Eriococcodes

63. Ovary smooth, 6-locular. — Asia........................... Subgenus Eriococcus section Nymphanthus

64*. Stigmas entire, connate — filaments thickened at top. — Asia to Australia........................................ Subgenus Eriococcus section Emblicastrum

64*. Stigmas entire or emarginate. — Asia......................... Subgenus Eriococcus section Scepasma

64*. Stigmas free, bifid..................................... 65 (Subgenus Eriococcus section Eriococcus )

65. Flowers in all leaf axils. — Asia.............................. Subgenus Eriococcus section Eriococcus

65. Pistillate flowers on leafy panicles at end of branchlets and staminate flowers closer to the base of branchlets without leaves (see Brunel 1987). — Asia............................ Subgenus Eriococcus section Eriococcus .......................... subsection Spiciferens

66. Branchlets (bi-)pinnatiform; sepals often biseriate; staminate nectar disc often 3 massive emarginate (or 6 separate) segments to absent, stamens may be inserted on a wide receptaculum. Pollen 3-(syn)colporate............................... 67 (Subgenus Gomphidium )

66. Branchlets pinnatiform; sepal whorls indistinct; staminate nectar disc entire or segmented................. 79

67. Branchlets bipinnatiform....................... 68

67. Branchlets pinnatiform........................ 70

68. Axes incrustate or hirsutulous with red hairs; stamens 2‒6; pollen clypeate. — West Indies.......................... Subgenus Xylophylla section Hemiphyllanthus

68. Axes not incrustate or hirsutulous, hairs usually white; stamens mainly 3‒5 (up to 20); pollen 3-(syn)colporate .......................................... 69

69. Inflorescences glomerules; pollen diverse, often 3-colporate or porate with diorate colpi (see Webster & Carpenter 2002),exine vermiculate to pilate;fruit conspicuously veined. — South America........................................... Subgenus Conami section Nothoclema

69. Inflorescences glomerules or panicles; pollen 3- or 4- syn- colporate with vermiculate/rugulate exine ( Lobreau-Callen et al. 1988); fruit smooth. — Southeast Asia , mostly New Guinea..... Subgenus Gomphidium section Nymania

70. Disc absent or rudimentary in both sexes......... 71

70. Disc entire or segmented in both sexes.......... 74

71. Sepals 6, biseriate, inner whorl petal-like, pistillate sepals leafy; stamens 3; ovary 3-locula. Calyx in fruit saccate. — New Caledonia........... Subgenus Gomphidium ..... section Gomphidium subsection Physoglochidion

71. Sepals 5 or 6, not distinctly biseriate; stamens (3–)5(–15), ovary 3–5-locular............................ 72

72. Sepals 5; stamens 5; ovary 4- or 5-locular. — New Caledonia...... Subgenus Gomphidium section Leptonema

72. Sepals 5 sometimes 6; stamens mostly (3–)5(–15); ovary 3-locular................................... 73

73. Disc rudimentary in both sexes; filaments shorter than anthers, inserted on a wide receptaculum. — Southeast Asia, New Caledonia................................ Subgenus Gomphidium section Adenoglochidion

73. Disc rudimentary or absent in both sexes; filaments longer than anthers, diverging from centre of receptaculum. — New Caledonia............. Subgenus Gomphidium . section Adenoglochidion subsection Eleutherogynium

74. Stamens connate............................ 75

74. Stamens free............................... 76

75. Inflorescences glomerules; pollen diverse, often 3-colporate or porate with diorate colpi (see Webster & Carpenter 2002), exine vermiculate to pilate; fruit conspicuously veined. — South America.................................... Subgenus Conami section Nothoclema

75. Inflorescences glomerules or paniculate; pollen 3- or 4- syncolporate, exine vermiculate/rugulate ( Lobreau-Callen et al. 1988); fruit smooth. — Southeast Asia, mainly New Guinea..... Subgenus Gomphidium section Nymania

76. Sepals 5, not distinctly biseriate; stamens mostly (3–) 5(‒15), filaments free; disc consisting of 3 emarginate segments or absent. — Southeast Asia, New Caledonia........ Subgenus Gomphidium section Adenoglochidion

76. Sepals 5 or 6, often biseriate (except in P. tuerckheimii G.L.Webster ); stamens 3, filaments free or connate; disc consisting of 3 emarginate segments or 6 free segments .......................................... 77

77. Sepals 5; pollen grains not syncolpate, colpi without distinct borders; exine reticulate. — South America............... Subgenus Gomphidium section Calodictyon

77. Sepals 6; pollen grains with marginate colpi, often meeting at poles; exine reticulate or ± vermiculate......... 78

78*. Inflorescences axillary cymules; pollen 3-syncolporate with fine to course reticulate exine. — Southeast Asia, New Caledonia Subgenus Gomphidium section Gomphidium

78*. Inflorescences glomerules or panicles; pollen 3-syncolpo- rate with vermiculate/rugulate exine. — Southeast Asia, mostly New Guinea....................................... Subgenus Gomphidium section Nymania

78*. Inflorescences glomerules; pollen diverse, often 3-colporate with diorate colpi (see Webster & Carpenter 2002), exine vermiculate to pilate. Fruit conspicuously veined. — South America... Subgenus Conami section................................... Nothoclema

79. Staminate disc entire. — South America...................... Subgenus Xylophylla section Adianthoides

79. Staminate disc segmented..................... 80

80. Filaments free or only fused at base............. 81

80. Filaments fused at least partially to completely, sometimes fused in separate sets or whorls................. 89

81. Leaves often with glands; anthers apiculate. Leaves thick; style entire. — South America.............................. Subgenus Emblica section Microglochidion

81. Leaves without glands; anthers not apiculate....... 82

82. Stamens 3................................. 83

82. Stamens 4 or 5.............................. 85

83. Brachyblasts often present; inflorescences cauliflorous; sepals 6. — South America................................... Subgenus Kirganelia section Ciccopsis

83. Brachyblasts absent; inflorescences axillary; sepals 5 84

84. Leaf blades <8 cm long; anther connective enlarged; pollen 4-colporate. — South America.. Subgenus Phyllanthus ......... section Phyllanthus subsection Clausseniani

84. Leaf blades> 8 cm long; anther connectives not enlarged; pollen perisyncolporate. Pistillate pedicel quite massive (up to 3 cm wide ( Brunel 1987)), fruit ornamented. — Africa....... Subgenus Ceramanthus section Ebolowani

85. Pistillate sepals 8–10. — South America....................... Subgenus Xylophylla section Diplocicca

85. Pistillate sepals 5 or 6......................... 86

86. Brachyblasts present......................... 87

86. Brachyblasts absent.......................... 88

87. Inflorescences (stalked) fascicles; stamens 5, filaments completely free; fruits 3-locular, dehiscent; seeds kidneyshaped, smooth with mottled patterns (similar to seeds of P. juglandifolius Willd. ). — Africa.......................... Subgenus Kirganelia section Omphacodopsis

87. Inflorescences panicles; stamens 4 or 5, filaments free or sometimes basally fused; fruits 3–5-locular, indehiscent; seeds globular, smooth. — Africa............................... Subgenus Kirganelia section Polyanthi

88. Stamens 3–5, free or slightly fused at base; anthers dehiscing vertically; fruits capsular; seeds smooth or faintly longitudinally striate. — Africa, Madagascar, Asia............................ Subgenus Anesonemoides

88. Stamen 2‒5(‒7); anthers dehiscing horizontally; fruits capsular; seeds smooth. — South America...................... Subgenus Emblica section Pityrocladus

89. Stamens fused in several whorls or sets.......... 90

89. Stamens fused in a central column.............. 93

90. Brachyblasts present; stamens fused in two sets with one central column and two separate free stamens..... 91

90. Brachyblasts absent; stamens in 2 or 3 whorls, fused in various ways. — West Indies............................. 94 (Subgenus Xylophylla section Willamia )

91. Pollen 3-colporate, exine pilate or reticulate....... 92

91. Pollen clypeate, exine areolate. — West Indies............... 94 (Subgenus Xylophylla section Willamia )

92. Exine pilate. — Africa...................................... Subgenus Kirganelia section Brazzeani

92. Exine reticulate. — Africa and Asia.......................... Subgenus Kirganelia section Anisonema

93. Stems and branchlets incrustate with dark platelets of bark or lenticellate...................................... 94 (Subgenus Xylophylla section Willamia )

93. Stems smooth.............................. 95

94*. Stems smooth;leaves alternate;stamens 3–15 in 3 whorls, connate in various ways; stigmas erect but not lacerate. Sepals 5 or 6. — West Indies... Subgenus Xylophylla ............ section Williamia subsection Discolores

94*. Stems and branchlets incrustate with small dark platelets on the fissured bark; leaves alternate; stamens (2–)3–6 with filaments connate, usually in 2 whorls; stigmas erect, conspicuously lacerate (see Webster 1958). — West Indies.............Subgenus Xylophylla section Williamia ........................... subsection Incrustati

94*. Stems smooth but prominently lenticellate; leaves opposite; stamens 5 with filaments connate, but 2 anthers inserted lower than the other 3; stigmas reflexed and cov- ering the ovary, apex sometimes blunt. — West Indies.............Subgenus Xylophylla section Williamia ............................. subsection Mirifici

95. Pollen 3–5-colporate or 5-brevicolporate ( Webster & Carpenter 2008), exine reticulate, microperforate or scabrous. — Africa and Asia................... 96

95. Pollen clypeate, exine areolate. — Americas.............................. 99 (Subgenus Xylophylla )

96. Anthers dehiscing obliquely to horizontally; exine microperforate or scabrous. — Pantropical, mostly African......................... Subgenus Afroswartziani

96. Anthers dehiscing vertically; exine reticulate................................. 97 (Subgenus Emblica )

97. Sepals mostly 5; staminate disc consisting of 5 segments; stamens 2‒5(‒7); anthers dehiscing horizontally; pollen 3‒5-colporate; pistillate disc entire or segmented. — South America Subgenus Emblica section Pityrocladus

97. Sepals 6; staminate disc consisting of 6 segments; stamens 3; anthers dehiscing vertically; pollen 4- or 5-colporate or 5-brevisulcate; pistillate disc entire.. 98

98. Inflorescences at terminal end of branchlet and secondary ramification on more basal end; pollen 5-brevisulcate. — Asia......... Subgenus Emblica section Botryoides

98. Inflorescences found along entire branchlet; pollen 4- or 5-colporate. — Asia Subgenus Emblica section Emblica

99. Inflorescences cauliflorous thyrses; stigmas petaloid. — West Indies.. Subgenus Xylophylla section Epistylium

99. Inflorescences axillary cymules; stigmas tapering, not petaloid, sometimes fused into a tube........... 100

100. Leaf blades mostly 1–2 cm long, with mesophyllar sclereids; stamens 3‒7. — West Indies.......................... Subgenus Xylophylla section Orbicularia

100. Leaf blades> 2 cm long, sometimes with mesophyllar sclereids; stamens 2‒7(‒8).................. 101

101. Brachyblasts often present; sepals 5; staminate disc consisting of 5 segments; stamens 3 (rarely 4); fruit a large fleshy capsule (usually> 2 cm diam). — West Indies........... Subgenus Xylophylla section Omphacodes

101. Brachyblasts absent; sepals 4–6; staminate disc usually consisting of 6 segments; stamens 2–7(–8); fruit small dry capsule (<1 cm diam).................... 102

102. Staminate sepals 5, pistillate sepals 6; inflorescences mostly unisexual cymules appearing after the leaves, several pistillate flowers per node; stamens 3‒7, thecae dehiscing horizontally; style present, elongated and ex- erted from calyx, stigmas dilated, bifid to multifid. — South America......Subgenus Xylophylla section Oxalistylis

102. Sepals in both sexes 4‒6; inflorescences bisexual cymules appearing with the expanding leaves on new branchlets, usually only 1 or 2 pistillate flowers among several staminate flowers; stamens 2‒6 (or 8), thecae dehiscing vertically; style like an erect tube, stigma branches nar- rowed to acute tips. — West Indies........................ Subgenus Xylophylla section Thamnocharis