Phyllantheae

Bouman, Roderick W., Kebler, Paul J. A., Telford, Ian R. H., Bruhl, Jeremy J., Strijk, Joeri S., Saunders, Richard M. K., Esser, Hans-Joachim, Falcón-Hidalgo, Banessa & Van, Peter C., 2022, A revised phylogenetic classification of tribe Phyllantheae (Phyllanthaceae), Phytotaxa 540 (1), pp. 1-100 : 7-9

. Key to the genera of tribe Phyllantheae

With the following key the genera of tribe Phyllantheae can be identified, but it does not take into account any exceptions or subgenera and (sub)sections. An asterisk (*) denotes parts of the key that are not dichotomous.

1. Pistillode present in staminate flowers ...............................................................................................................................................2

1. Pistillode absent in staminate flowers ................................................................................................................................................5

2. Stamens> 10—Africa & Madagascar........................................................................................................................... Lingelsheimia

2. Stamens 4–7........................................................................................................................................................................................3

3. Branching non-phyllanthoid (laminate leaves and flowers on all axes, branchlets not deciduous); staminate sepals 4‒7 (usually 5), stamens 4‒7, filaments free or fused for half of length; anther connectives non-apiculate; fruits capsules or baccate.....................4

3. Branching phyllanthoid (leaves on main stem reduced to scales, = cataphylls, laminate leaves and flowers on lateral axes, lateral branchlets deciduous); staminate sepals 4, stamens 4, filaments connate, anther connectives apiculate; fruits capsules—Asia........ ............................................................................................................................................................................. Glochidion ( G. moi )

4. Petals absent; fruits with 2 seeds per locule; pollen exine verruculose—Pantropical .......................................................... Flueggea

4. Petals present; fruits with 1 or 2 seeds per locule; pollen exine reticulate—West Indies ................................................ Heterosavia

5. Disc absent (sepal scales may be present, these close flower when anthers are unripe, no glandular function) ...............................6

5. Disc / disc glands present .................................................................................................................................................................10

6. Inflorescences on specialized leafless axes; staminate sepals 4, stamens (2‒)3‒4(‒5), filaments free; fruits indehiscent, woody or drupaceous............................................................................................................................................subgenus Cicca section Cicca

6. Inflorescences in axils of leaves to cauliflorous; staminate sepals usually 5 or 6, stamens 2‒15, filaments free or connate; fruits capsules...............................................................................................................................................................................................7

7. Staminate flowers without sepal scales; filaments free or connate, anther connectives sometimes apiculate; stigmas usually entire or erect and tightly together; ovary 3‒15-locular ...............................................................................................................................8

7. Staminate flowers often with sepal scales; filaments connate, anther connectives not apiculate; stigmas usually bifid; ovary 3- locular.................................................................................................................................................................................................9

8. Staminate sepals spreading or united in a tubular shape; disc lobes present; filaments free; ovary 3‒5-locular— Malesia................ ............................................................................................................................................................................... Dendrophyllanthus

8. Staminate sepals often recurved; disc absent; filaments tightly together (separating as flowers age); ovary 3‒15-locular— Australia, mainland Asia, Malesia, Pacific ......................................................................................................................................... Glochidion

9. Sepal scales often present in staminate flowers; fruits wider than long; seeds smooth .......................................................... Breynia

9. Sepal scales absent in staminate flowers (except in Synostemon bacciformis ); fruits longer than wide; seeds sculptured................. ........................................................................................................................................................................................... Synostemon

10. Branching non-phyllanthoid or sub-phyllanthoid (leaves at base of branchlets not reduced to scales (often in juveniles), lateral branchlets deciduous) .......................................................................................................................................................................11

10. Branching phyllanthoid ....................................................................................................................................................................19

11. Stamens> 10—Africa & Madagascar........................................................................................................................... Lingelsheimia

11. Stamens 2‒5......................................................................................................................................................................................12

12. Sepals 4; staminate disc entire; stamens 4, filaments free; seeds with blue sarcotesta—Pantropical............................ Margaritaria

12. Sepals 5 or 6 (rarely 4, but then with 2 stamens); staminate disc entire or segmented; stamens 2 or 3, filaments free or connate; seeds with no or whitish sarcotesta ..................................................................................................................................................13

13. Staminate disc entire (H-shaped in Moeroris arenaria )...................................................................................................................14

13. Staminate disc segmented into glands..............................................................................................................................................15

14*. Sepals 6 in both sexes; staminate disc urceolate; stamens 3, filaments connate—Asia........................................................ Cathetus

14*. Sepals 6 in both sexes; staminate disc not urceolate; stamens 2, filaments free—Africa............................................... Plagiocladus

14*. Sepals 4 in staminate flowers, 6 in pistillate flowers; staminate disc H-shaped around filaments; stamens 2, filaments free—North America .......................................................................................................................................................... Moeroris ( M. arenaria )

15. Sepals 5; stamens 5, filaments free—Africa ............................................. Kirganelia subgenus Kirganelia section Pseudomenarda

15. Sepals 5 or 6; stamens usually 3 (rarely 2), filaments connate (free in Moeroris rosmarinifolius )—Africa to Asia.......................16

16. Plagiotropic branchlets caducous,usually fascicled (non-phyllanthoid in section Elutanthos , but then with paniculate inflorescences); fruits capsules or berries.................................................................................................................................................... Phyllanthus

16. Plagiotropic branchlets persistent; inflorescences axillary fascicles, never paniculate; fruits capsules ..........................................17

17. Leaves on all axes spirally arranged; filaments connate; pistillate disc consisting of free glands—(North America, Africa and Asia/ Australia) .................................................................................................................................................................................. Nellica

17. Leaves on all axes distichous (except at basal nodes in Cicca ); filaments free or connate; pistillate disc mostly entire (when segmented then filaments mostly free and leaves always distichous)..............................................................................................18

18*. Leaves distichous; pollen 4-colporate; seeds smooth or verrucate—Americas ................................................................................... ..................................................................................................................... Phyllanthus subgenus Phyllanthus section Loxopodium

18*. Leaves distichous; pollen clypeate; seeds smooth or verrucate—Africa, Asia, Australia and Pacific... Cathetus subgenus Macraea

18*. 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................................................................................. Cicca subgenus Betsileani

19. Herbs or subshrubs (small plants with woody base) ........................................................................................................................20

19. Shrubs to trees, rarely climbers ........................................................................................................................................................25

20. Inflorescences unisexual...................................................................................................................................................................21

20. Inflorescences bisexual.....................................................................................................................................................................24

21. Pistillate inflorescences on proximal position and staminate inflorescences on distal position on plagiotropic branchlets; seeds transversely ribbed—ovary often covered with tubercles—pantropical, but origin Asia .......................... Emblica section Urinaria

21. Pistillate inflorescences on distal position and staminate inflorescences on proximal position on plagiotropic branchlets; seeds smooth or longitudinally or transversely striate, but not ribbed ......................................................................................................22

22. Cataphyllary stipules not auriculate .................................................................................................................................. Phyllanthus

22. Cataphyllary stipules (unilaterally) auriculate..................................................................................................................................23

23. Filaments connate, anthers free; seeds smooth or longitudinally striate—Africa................................................................. Moeroris

23. Filaments usually free, sometimes connate, anthers free or connate in a synandrium; seeds smooth, striate or verrucate—South America ............................................................................................................................................................................. Phyllanthus

24*. Stamens 3, filaments free, anthers dehiscing vertically; pistillate disc entire; pollen 3-syncolporate, exine reticulate; stigmas entire or emarginate; seeds smooth or with faintly striate— Australia, Malesia, Pacific ................. Dendrophyllanthus section Leptonema

24*. Stamens 2 or 3, filaments entirely or partially connate (free in M. arenaria ), anthers dehiscing oblique to horizontally (vertically in M. arenaria ); pistillate disc entire; pollen 3-colporate, exine reticulate; seeds smooth or longitudinally striate—North America, pantropical invasive............................................................................................................................. Moeroris subgenus Swartziani

24*. Stamens 3, filaments free, anthers dehiscing horizontally; pistillate disc segmented; pollen grains brevicolporate and diorate or porate, exine pilate; seeds verruculose—South America ....................................... Phyllanthus subgenus Conami section Apolepsis

25. Sepals 4 in staminate flowers; stamens 2, filaments connate, anthers dehiscing horizontally ( Nymphanthus or Phyllanthus chryseus ) or vertically ( Phyllanthus section Thamnocaris )—pollen pantoporate or clypeate.........................................................................26

25. Sepals 5 or 6 in staminate flowers (4 in Cicca acida , but then stamens 4, filaments free); stamens 3–15, filaments free or connate, anthers dehiscing mostly vertically, sometimes horizontally ...........................................................................................................27

26. Inflorescences mostly unisexual; staminate disc segmented; anthers dehiscing horizontally (except in N. ruber & N. touranensis ); pollen pantoporate—Asia............................................................................................................................................... Nymphanthus

26. Inflorescences unisexual ( P. chryseus ) or bisexual; staminate disc entire ( P. chryseus ) or segmented; anthers dehiscing horizontally ( P. chryseus ) or vertically (P. section Thamnocaris ); pollen clypeate—South America................................................... Phyllanthus

27. Filaments fused in sets, rarely free ( K. glauca & K. flexuosa ), stamens 2 ( K. flexuosa ) or 5, connectives not apiculate; fruits baccate—Africa to Australia.............................................................................................................................................. Kirganelia

27. Filaments never in sets, free or connate, stamens 3‒20, connectives sometimes apiculate; fruits capsules or baccate (then with 3 connate stamens) ..............................................................................................................................................................................28

28. Plagiotropic branchlets sometimes differentiated in vegetative (with larger leaves) and floriferous (with smaller leaves) branchlets, pinnatiform; stamens 3, filaments connate, anthers dehiscing vertically, connectives apiculate; pistillate disc segmented—Asia & Pacific................................................................................................................................................................................. Glochidion

28. Plagiotropic branchlets not differentiated and all with leaves of similar size (or unifoliate), sometimes bipinnatiform; stamens 3–20, filaments connate or free, anthers dehiscing horizontally to vertically, connectives apiculate or not; pistillate disc entire..29

29. Branchlets (bi-)pinnatiform; sepals often biseriate (inner whorl much longer); staminate disc often of 3 massive emarginate (or 5 separate) segments to absent, stamens may be inserted on a wide receptaculum; stamens usually 3 or 5 (up to 20 in Pacific species); stigmas mostly entire, rarely bifid—pollen 3–(syn-)colporate.........................................................................................................30

29. Branchlets pinnatiform; sepal whorls indistinct; staminate disc segmented or entire; stamens 2–7(– 15 in species of South America and West Indies) stigmas usually bifid or lacerate ...........................................................................................................................31

30. Inflorescences fascicles or panicles; sepals 5 or 6, filaments free or connate, anthers dehiscing vertically to obliquely, connectives usually apiculate; pollen 3- or 4-syncolporate, exine rugulose-reticulate, vermiculate, pilate or ± vermiculate—Calyx in fruit sometimes saccate— Malesia, Australia, Pacific ................................................................................................... Dendrophyllanthus

30. Inflorescences fascicles; sepals 6 (5 in P. tuerckheimii ), filaments connate (free in P. tuerckheimii ), anthers dehiscing horizontally (vertically in in P. tuerckheimii ), connectives rarely elongated; pollen diverse, often 3-colporate or porate with diorate colpi (see Webster & Carpenter 2002), exine vermiculate to pilate—Fruits conspicuously veined—South and Central America..................... ................................................................................................................................... Phyllanthus subgenus Conami section Conami

31*. Branchlets never transformed to phylloclades; sepals 6; staminate disc segmented; stamens 3, filaments connate, anthers dehiscing vertically, connectives often apiculate; fruits capsules (drupe in E. officinalis )—Asia......................................................... Emblica

31*. Branchlets never transformed to phylloclades; sepals 5 or 6; staminate disc segmented; stamens 3, filaments connate, anthers dehiscing mostly horizontally to obliquely, connectives rarely apiculate; fruits capsules—mainly Africa.......................... Moeroris

31*. Branchlets sometimes transformed to phylloclades; sepals 4‒8; staminate disc mostly segmented, sometimes entire; stamens 3‒15, filaments free or fused, sometimes fused in several whorls, anthers dehiscing horizontally to vertically, connectives sometimes apiculate; fruits capsules or berries—Americas............................................................................................................… Phyllanthus