Abrahamia Randrian. & Lowry, 2017

Randrianasolo, Armand, Lowry II, Porter P. & Schatz, George E., 2017, Taxonomic treatment of Abrahamia Randrian. & Lowry, a new genus of Anacardiaceae from Madagascar, Boissiera 71, pp. 1-152 : 13-30

publication ID

https://doi.org/ 10.5281/zenodo.7618113

DOI

https://doi.org/10.5281/zenodo.7618382

persistent identifier

https://treatment.plazi.org/id/038887C6-FFC7-FFCB-FE43-86171BEB0FF3

treatment provided by

Carolina

scientific name

Abrahamia Randrian. & Lowry
status

gen. nov.

Abrahamia Randrian. & Lowry View in CoL View at ENA , gen. nov.

Typus: Abrahamia ditimena (H. Perrier) Randrian. & Lowry View in CoL (≡ Protorhus ditimena H. Perrier View in CoL ).

Diagnosis

Dioecious trees or shrubs. Leaves simple, persistent, opposite, subopposite, verticillate or alternate, chartaceous, subcoriaceous or coriaceous, lateral veins parallel, arcuate, forming an obtuse to acute angle with the midrib toward the base of the blade. Inflorescence erect, a thyrse or panicle. Flowers (4-)5(-6)-merous; pedicel short, articulated; sepals much smaller than petals; stamens alternipetalous, filament broadened at the base (sterile in female flowers, forming staminodes), anthers dorsifixed, introrse, glabrous; ovary pubescent or glabrous (rudimentary or absent in male flowers), with 1 locule containing a single subpendulous or pendulous ovule. Fruits drupaceous, ellipsoidal or ovoid, symmetrical or slightly asymmetrical, resiniferous; seeds ruminate; cotyledons usually inseparable (except in Abrahamia ibityensis (H. Perrier) Randrian. & Lowry , in which the resiniferous canals are only peripheral).

Description

Dioecious trees or shrubs; bark with white, translucent or red resinous latex. Leaves alternate, opposite or subopposite (verticillate in one species), simple, coriaceous, subcoriaceous or chartaceous, persistent (deciduous in A. humbertii ). Venation pinnate, craspedodromous or semicraspedromous, midvein usually flat, rarely very slightly raised above, prominently raised below (rarely on both surfaces), secondary veins parallel or nearly so, straight or arcuate, prominent below or on both surfaces, forming an obtuse to acute angle with the midrib toward the base of the blade. Inflorescence terminal and/ or axillary, an erect thyrse or panicle, or intermediate and forming a paniculiform thyrse. Flowers usually 5-merous (sometimes 4- or 6-merous); sepals much smaller than petals; stamens 5 or 4 (rarely 6), alternipetalous, filaments straight, flattened, broadened at the base, inserted basally on the outer surface of the disk, anthers dorsifixed, introrse, dehiscent with longitudinal slits (stamens reduced in size and sterile in female flowers, forming staminodes); disk usually cupuliform and crenulate in male flowers, annular and cupuliform in female flowers; ovary with 1 locule containing a single pendulous or subpendulous or ovule on a short funiculus, placentation subapical (ovary rudimentary or absent in male flowers); style one, short, apically trifid or unbranched with 3 capitate stigmatic lobes. Fruits drupaceous, ellipsoidal or ovoid, symmetrical or slightly asymmetrical, exocarp usually with longitudinal striations on surface, glabrous or velutinous, mesocarp resinous, endocarp thin, papyraceous; seed solitary, with resiniferous canals throughout (only peripheral in A. ibityensis ), ruminate; cotyledons usually inseparable (separable in A. ibityensis ).

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Etymology

The genus is named in honor of the late Jean Prosper Abraham (1936-1996), a wellknown Malagasy forester with a remarkable knowledge of the woody plants of Madagascar and who contributed much to our knowledge of the island’s rich and diverse flora.

Notes

Abrahamia includes a total of 34 species, all endemic to Madagascar, including 15 species formerly placed in Protorhus (e.g., by PERRIER DE LA BâTHIE, 1944, 1946), to which 19 additional species are newly described here. With the recognition of Abrahamia as a separate genus, Protorhus is now restricted to a single species in Africa, P. longifolia . The two genera differ in several flower and fruit characters, as indicated by PELL (2004) and PELL et al. (2011), and summarized in Table 1 View Table 1 .

Abrahamia is represented in all of Madagascar’s major vegetation types (MOAT & SMITH, 2007) and broad bio-climatic zones ( CORNET, 1974; SCHATZ, 2000). Species can be found in humid forest in the eastern and Sambirano regions ( A. antongilensis , A. sambiranensis ), wooded grassland and bushland in the center of the island ( A. ibityensis , A. buxifolia ), western dry forest ( A. deflexa , A. humbertii ), and southwestern subarid spiny forest-thicket ( A. phillipsonii Randrian. & Lowry ). Members of the genus occur on many types of substrate and at a wide range of elevations, from littoral forest on sand at the sea level ( A. nitida (Engl.) Randrian. & Lowry , A. suarezensis Randrian. & Lowry ) to humid forest on lateritic soil at altitudes up to 2000 m ( A. buxifolia , A. itremoensis ).

Species of Abrahamia were described by PERRIER DE LA BâTHIE (1946) as polygamo-dioecious, indicating that he interpreted them as having both unisexual and hermaphroditic flowers on the same individual. However, careful observation has shown that all members of the genus are functionally dioecious. Female flowers have rudimentary, sterile stamens that are reduced in size, whereas male flowers either have an ovary that is rudimentary and non-functional or lack an ovary altogether.

16 Boissiera 71

Key to the species of Abrahamia View in CoL

[Numbers in parentheses indicate the previous coupLet Leading to the present one]

1. Leaves completely glabrous on both surfaces (occasionally with extremely small indument visible only under a dissecting microscope on or adjacent to the midvein on the abaxial surface)...................................................................................................... 2

1a. Leaves with at least a few evident trichomes, sometimes restricted to the midvein beneath ..................................................................................................................... 20

2. Leaves usually verticillate (3 per node, sometimes one leaf broken off but leaving a scar); E coast littoral forest (occasionally to 200 m elevation).............................. 23. A. nitida View in CoL

2a. Leaves alternate or opposite to subopposite................................................................. 3

3. Largest leaf blade at least 15 cm long........................................................................... 4

3a. Largest leaf blade less than 15 cm long (rarely to 17 cm in A. oblongifolia View in CoL and A. littoralis View in CoL ) .................................................................................................................. 6

4. Petiole stout, c. 4-5 mm in diam.; young branches with corky bark; littoral and low elevation humid forest, Maroantsetra S to Tampolo ................................ 15. A. latifolia View in CoL

4a. Petiole slender, no more than 2(-2.5) mm in diam.; young branches without corky bark; humid forest................................................................................................................ 5

5. Largest leaf with c. 50-60 pairs of secondary veins; fruits ellipsoid, covered with dense, short indument; Betampona .......................................................... 2. A. betamponensis View in CoL

5a. Largest leaf with c. 30-45 pairs of secondary veins; fruits obovoid, glabrous; Lokobe and Tsaratanana ........................................................................................19. A. lokobensis View in CoL

6 (3). Leaves elliptic to ovate (rarely a few slightly obovate), broadest at the middle or in the proximal part of the blade ........................................................................................... 7

6a. Leaves obovate to oblanceolate, nearly all broadest in the distal part of the blade......... 11

7. Leaves narrowly elliptic to narrowly ovate (rarely a few slightly obovate), at least (2.5-) 3 times as long as wide; dry forest, Isalo to Andohahela ....................27. A. phillipsonii View in CoL

7a. Leaves elliptic to ovate, sometimes broadly so (rarely a few slightly obovate), no more than 1.5-2 times as long as wide.................................................................................. 8

8. Fruits covered with dense short indument; flowers 4-merous (rarely 5-merous); humid forest, Marojejy and Anjanaharibe-Sud S to Ranomafana and Ivohibe............................ ..........................................................................................................17. A. lenticellata View in CoL

8a. Fruits glabrous; flowers 5-merous (unknown in A. darainensis View in CoL ) ................................... 9

9 (8). Leaves orbicular to broadly elliptic (rarely a few broadly ovate), blades no more than 2 times as long as wide, subcoriaceous, apex rounded (rarely broadly acute); dry forest, calcareous substrates, Ankarana, Bemaraha .........................................12. A. humbertii View in CoL

9a. Leaves elliptic to ovate (occasionally a few broadly ovate or obovate), most blades at least 2 times as long as wide, chartaceous, apex acute to acuminate .......................... 10

10. Blade of largest leaf to c. 6.5 cm long, often discolorous, the upper surface dark green to chocolate brown, apex emarginate or rounded to acute; dry western forest (Bemaraha, Ketsepy, Antsalova, Marovoay, Mahajanga) ................................................ 6. A. deflexa View in CoL

10a. Blade of largest leaf c. 8 cm long, concolorous, the upper surface drying grayish green, apex acuminate; humid forest, Daraina............................................... 5. A. darainensis View in CoL

11 (6). Largest leaf with 40-70 pairs of secondary veins; humid eastern forest ...... 30 A. sericea View in CoL

11a. Largest leaf with 10-40 pairs of secondary veins ........................................................ 12

12. Inflorescences all axillary; leaves alternate; E coast littoral forest, Tampolo S to Mahabo ............................................................................................................... 18. A. littoralis View in CoL

12a. Inflorescences terminal and sometimes also axillary; leaves opposite to subopposite........13

13. Tertiary venation obscure on abaxial surface ............................................................. 14

13a. Tertiary venation visible on abaxial surface, at least toward the margins....................16

14. Secondary veins closely spaced, 6-9 pairs per cm in median portion of largest leaf, blade distinctly glaucous on adaxial surface, nearly always strongly discolorous when dry, often chocolate brown above (sometimes obscured by glaucescence); fruit velutinous (sometimes glabrescent); dry forest, central high plateau, Itremo, Ibity, Ambatofinandrahana ............................................................................................................................3. A. buxifolia View in CoL

14a. Secondary veins widely spaced, 3-5 pairs per cm in median portion of largest leaf, blade neither glaucous (sometimes weakly so in A. suarezensis View in CoL ) nor strongly discolorous when dry, various shades of green but not chocolate brown above; fruit glabrous............... 15

15. Largest leaf with 15-30 pairs of secondary veins, margins distinctly revolute; trees 15-20 m tall; littoral and dry forest, Mt. d’Ambre, S of Vohémar and Masoala.......................... .............................................................................................................. 28. A. revoluta View in CoL

15a. Largest leaf with 12-15 pairs of secondary veins, margins slightly thickened but not revolute; trees to 10(-12) m tall; N dry and coastal forests, Antsiranana, Vohémar, Daraina............................................................................................. 31. A. suarezensis View in CoL

16 (13).Leaves with venation craspedodromous, secondary veins all clearly reaching the margin (rarely a few terminating away from the margin in A. oblongifolia View in CoL ).............................. 17

16a. Leaves with venation semicraspedodromous, at least some secondary veins branching and not reaching the margin...................................................................................... 19

18 Boissiera 71

17 (16).Largest leaf with 40-70 pairs of secondary veins, largest blade 9-15 cm long ..... ................................................................................................................ 30. A. sericea View in CoL

17a. Largest leaf with 15-33 pairs of secondary veins, largest blade 2.5-7.5(-8.5) cm long.... .................................................................................................................................. 18

18. Blade of largest leaf at least (4.5-) 5 cm long, coriaceous, rigid; mature fruit 2-3 × 1.4-1.6 cm ............................................................................................................... 8. A. ditimena View in CoL

18a. Blade of largest leaf no more than 4-(4.5) cm long, subcoriaceous, not rigid; mature fruit 1-1.5 × 0.5-0.8 cm ............................................................................21. A. minutifolia View in CoL

19 (16).Largest leaf blade 8-14(-17) cm long, petiole c. 2 mm in diam.; fruits glabrous; dry to humid forest, Nossi Be, Antsiranana area .......................................... 24. A. oblongifolia View in CoL

19a. Largest leaf blade 3.5-6 cm long, petiole c. 0.5-1 mm in diam.; fruits with short dense indument; littoral forest and low-elevation humid forest, Tolagnaro area ...................... ......................................................................................................... 7. A. delphinensis View in CoL

20 (1). Blade of largest leaf at least 10 cm long...................................................................... 21

20a. Blade of largest leaf no more than 10 cm long .......................................................... 34

21. Largest leaves with at least 40 pairs of secondary veins..............................................22

21a. Largest leaves with fewer than 40 pairs of secondary veins ........................................26

22. Inflorescences all axillary...........................................................................................23

22a. Inflorescences terminal and sometimes also axillary ..................................................24

23. Leaves with evident tertiary venation, apex flat; fruits glabrous; humid forest, Marojejy, Anjanaharibe-Sud, Masoala, Rantabe, Ambalabe .................................. 10. A. elongata View in CoL

23a. Leaves lacking evident tertiary venation, apex usually folded in pressed material; fruits pubescent when young, glabrescent at maturity; humid forest, Ambilobe, Ampondrabe, Ambato, Kalabenono-Galoka, Ambanja, Andavakoera ................ 29. A. sambiranensis View in CoL

24 (22).Secondary veins in median portion of leaf blade closely spaced, c. 5-6 per cm; fruit with dense indument; humid eastern forest..................................................... 30. A. sericea View in CoL

24a. Secondary veins in median portion of leaf blade more widely spaced, c. 3 per cm; fruit glabrous or with evident indument............................................................................25

25. Leaves with an acute to short acuminate apex (rarely a few rounded); fruits with dense, short indument; flowers 4-merous (rarely 5-merous); humid forest............................... ......................................................................................................... 17. A. lenticellata View in CoL

25a. Leaves with a broadly acute to rounded or retuse apex; fruits glabrous; flowers 5-merous; dry forest...........................................................................................11. A. grandidieri View in CoL

26 (21).Inflorescences all axillary...........................................................................................27

26a. Inflorescences terminal and sometimes also axillary ..................................................29

27. Abaxial surface of leaves with dense, erect to spreading light brown lanate indument ... .............................................................................................................. 34. A. viguieri View in CoL

27a. Abaxial surface of leaves glabrous or with spare, short indument ..............................28

28. Longest petiole 5-10 mm; eastern humid, low to mid-elevation forest... 16. A. lecomtei View in CoL

28a. Longest petiole (18-) 20-30 mm; E coast littoral forest, Tampolo S to Mahabo ............... ..............................................................................................................18. A. littoralis View in CoL

29 (26).Leaf apex long acuminate to nearly caudate, blade chartaceous .................................30

29a. Leaf apex acute to retuse or emarginate, sometimes short-acuminate (acumen <1 cm long), blade chartaceous to subcoriaceous or coriaceous .......................................... 32

30. Abaxial surface of leaves densely covered with reddish-brown lanate indument, venation craspedodromous; eastern littoral forest to premontane forest on sand or laterite, Maroantsetra, Toamasina, Zahamena PN, Fort-Carnot ........................... 34. A. viguieri View in CoL

30a. Abaxial surface of leaves glabrous or with scattered, white sericeous indument, venation semi-craspedodromous .............................................................................................31

31. Adaxial surface of leaves always glabrous, widest blade to 3.5(-4) cm; fruit velutinous; eastern humid forest .........................................................................17. A. lenticellata View in CoL

31a. Adaxial surface of leaves glabrous or with scattered white appressed indument, widest blade at least 4 cm; fruit glabrous; north-eastern humid forest, Masoala, Maroantsetra .........................................................................................................1. A. antongilensis View in CoL

32 (29).Leaves with 10-20 pairs of widely spaced secondary veins, only 1 or 2 per cm in median portion of blade; eastern and north-eastern humid forest, Maroantsetra, Manongarivo, Andapa, Sambava, Ranomafana, Mantadia, Zahamena ..................... 32. A. thouvenotii View in CoL

32a. Leaves with 20-40 pairs of narrowly spaced veins, 2-6 per cm in median portion of blade ........................................................................................................................33

33. Leaves obovate, never more than 2.5 times as long as wide, coriaceous; inflorescences terminal and sometimes also axillary; fruits covered with dense indument; humid eastern forest .......................................................................................... 8. A. ditimena View in CoL

33a. Leaves oblanceolate (sometimes narrowly elliptic), at least 3 times as long as wide, chartaceous to coriaceous, flexible; terminal and axillary; fruits glabrous; along rivers and streams, western and southern Madagascar, Ambanja, Befandriana Nord, Ambalavao, Ihosy, Sakaraha, Amboasary, Ampanihy .............................................11. A. grandidieri View in CoL

34 (20).Largest leaves with at least 40 pairs of secondary veins..............................................35

34a. Largest leaves with fewer than 40 pairs of secondary veins ........................................36

20 Boissiera 71

35 (34).Secondary veins in median portion of leaf blade closely spaced, c. 5-6 per cm; fruit with dense indument; humid eastern forest..................................................... 30. A. sericea View in CoL

35a. Secondary veins in median portion of leaf blade more widely spaced, c. 3 per cm; fruit glabrous; along rivers and streams, western and southern Madagascar, Ambanja, Befandriana Nord, Ambalavao, Ihosy, Sakaraha, Amboasary, Ampanihy ........................ ......................................................................................................... 11. A. grandidieri View in CoL

36 (34).Largest leaves with no more than 20 pairs of secondary veins....................................37

36a. Largest leaves with more than 20 pairs of secondary veins.........................................46

37. Leaves elliptic to orbicular or slightly ovate, broadest at or below the middle............38

37a. Leaves obovate to oblanceolate, broadest beyond the middle.....................................40

38. Leaves orbicular to broadly elliptic (rarely a few broadly ovate), no more than 2 times as long as wide; dry forest, Ankarana, Bemaraha .....................................12. A. humbertii View in CoL

38a. Leaves elliptic to narrowly elliptic, at least 2 times as long as wide ............................39

39. Largest leaves 2.5-3.6 cm wide; humid and transition forest, Manambato, Daraina, Androranga River valley near Sambava.................................................. 4. A. capuronii View in CoL

39a. Largest leaves 1.5-2(-2.2) cm wide; dry forest, Isalo to Andohahela............................... ......................................................................................................... 27. A. phillipsonii View in CoL

40 (37).Inflorescences all axillary, female inflorescences with no more than 5 flowers (male inflorescences unknown); humid forest, Nosy Be, Ampasindava, Ambongo-Boina ........ ........................................................................................................... 26. A. pauciflora View in CoL

40a. Inflorescences terminal and sometimes also axillary, with 8 to numerous flowers (rarely only 5 in female inflorescences of A. suarezensis View in CoL ) .......................................................41

41. Blade of largest leaf 2-3(-4) cm long; fruits 1-1.5 cm long .................22. A. minutifolia View in CoL

41a. Blade of largest leaf at least (3.5-) 4 cm long; fruits 1.5-2.5 cm long...........................42

42. Fruit densely covered with short indument; secondary veins semicraspedodromous, at least some branching and not reaching the margin; littoral forest, Tolagnaro area....... ......................................................................................................... 7. A. delphinensis View in CoL

42a. Fruit glabrous; secondary veins craspedodromous, all clearly reaching the margin....43

43. Petiole stout, (1.5-) 2-3 mm in diam. on largest leaves, tertiary venation distinctly raised on abaxial surface; eastern and north-western humid forest, Ambanja, Marojejy, Anajanaharibe Sud, Ranomafana, Analamazaotra, Zahamena ........... 32. A. thouvenotii View in CoL

43a. Petiole slender, no more than 1.5 mm in diam. on largest leaves, tertiary venation obscure to weakly raised on abaxial surface...............................................................44

44 (43).Secondary veins closely spaced, 5-6 pairs per cm in median portion of largest leaf; humid eastern forest, Analamazaotra, Ambalabe .................................... 21. A. louvelii View in CoL

44a. Secondary veins widely spaced, 3-4 pairs per cm in median portion of largest leaf....45

45. Inflorescences 1-3(-3.5) cm long; leaf margins undulate, secondary veins c. 10-12(-14) pairs; northern dry and coastal forest, Antsiranana, Vohémar, Daraina .......................... ........................................................................................................ 31. A. suarezensis View in CoL

45a. Inflorescences 4-9 cm long; leaf margins plane or only weakly undulate, secondary veins 14 or more pairs; humid forest, Ankazobe, E of Fianarantsoa, Ranomafana, Andohahela (Parcel #1), Ivohibe ................................................................................... 33. A. turkii View in CoL

46 (36).Venation semicraspedodromous, at least some secondary veins branching and not reaching the margin; fruit with evident indument ............................ 17. A. lenticellata View in CoL

46a. Venation craspedodromous, secondary veins all clearly reaching the margin; fruit glabrous or with evident indument .......................................................................... 47

47. Petiole of the largest leaves at least 10 mm long ....................................................... 48

47a. Petiole of the largest leaves less than 10 mm long .................................................... 53

48. Inflorescences all axillary; east coast littoral forest, Tampolo S to Mahabo ..................... ............................................................................................................. 18. A. littoralis View in CoL

48a. Inflorescences terminal and sometimes also axillary ................................................. 49

49. Tertiary veins of leaves barely visible, almost appearing absent; corolla of female flowers glabrous on outer surface ............................................................. 20. A. longipetiolata View in CoL

49a. Tertiary veins of leaves visible at least on one surface; corolla of female flowers with indument on outer surface ....................................................................................... 50

50. Leaves no more than 2.5 times as long as wide; fruit with evident indument ........... 51

50a. Leaves at least 3 times as long as wide; fruits glabrous ............................................. 52

51. Leaves chartaceous; littoral forest, Tolagnaro area............................. 7. A. delphinensis View in CoL

51a. Leaves coriaceous; humid eastern forest ................................................. 8. A. ditimena View in CoL

52 (50).Leaves obovate to elliptic; secondary veins 18-33 pairs, 1-4 mm apart; NE humid and sub-humid transitional forest (Daraina, Vohémar) ................................. 4. A. capuronii View in CoL

52a. Leaves oblanceolate; secondary veins 35-40-(47) pairs, 0.5-5 mm apart; dry and subhumid forest, W and SW.................................................................. 11. A. grandidieri View in CoL

53 (47).Secondary venation weakly raised on upper leaf surface; fruits glabrous .................. 54

53a. Secondary venation distinctly raised on upper leaf surface; fruits with evident indument ................................................................................................................. 60

22 Boissiera 71

54 (53).Fruits 3-3.5 cm long, ellipsoid; infructescence axes stout, 1.5-2 mm in diam.; largest leaves 8-9 × 2.5 cm, apex acute; littoral forest, Tolagnaro area............ 9. A. ellipticarpa View in CoL

54a. Fruits 1.5-2.6 cm long, ovoid or ellipsoid; infructescence axes slender, to c. 1(-1.2) mm in diam.; largest leaves either smaller than 8 × 2 cm or with a rounded or emarginate apex ..........................................................................................................................55

55. Leaves obovate to oblanceolate..................................................................................56

55a. Leaves elliptic to oblong, rarely a few slightly ovate or obovate..................................59

56. Largest leaves with at least 35 pairs of secondary veins; dry forest .... 11. A. grandidieri View in CoL

56a. Largest leaves with no more than 25(-28) pairs of secondary veins; humid forest......57

57. Secondary veins widely spaced, 3-4 pairs per cm in median portion of largest leaf blade; humid forest, Ankazobe, E of Fianarantsoa, Ranomafana, Andohahela (Parcel #1), Ivohibe .............................................................................................. 33. A. turkii View in CoL

57a. Secondary veins closely spaced, 5-6 pairs per cm in median portion of largest leaf blade ..................................................................................................................................58

58. Blade of largest leaf to 4(-4.2) cm long; Analamazaotra, Ambalabe; humid forest .......... ............................................................................................................... 21. A. louvelii View in CoL

58a. Blade of largest leaf at least (4.8-) 5 cm long; semi-deciduous and evergreen humid forest, Daraina .................................................................................................25. A. patrickii View in CoL

59 (55).Petiole slender, <1 mm in diam.; cotyledons fused; trees c. 3-12 m tall; dry forest, Isalo to Andohahela ................................................................................. 27. A. phillipsonii View in CoL

59a. Petiole stout, 1.2-1.5 mm in diam.; cotyledons separable; shrubs to small trees <2 m tall; sclerophyllous forest, Ibity............................................................ 13. A. ibityensis View in CoL

60 (53).Leaves glaucous on adaxial surface, nearly always strongly discolorous when dry, often chocolate brown adaxially (sometimes obscured by glaucescence); dry forest, central high plateau, Itremo, Ibity, Ambatofinandrahana.................................... 3. A. buxifolia View in CoL

60a. Leaves not glaucous on adaxial surface, uniform in color or weakly discolorous........61

61. Leaves puberulous on adaxial surface; Itremo area ........................... 14. A. itremoensis View in CoL

61a. Leaves glabrous on adaxial surface.............................................................................62

62. Leaves chartaceous; littoral forest, Tolagnaro area..............................7. A. delphinensis View in CoL

62a. Leaves coriaceous; eastern humid forest ................................................. 8. A. ditimena View in CoL

Clé des espèces d’Abrahamia

[Les chiffres entre parenthèses renvoient au coupLet de provenance]

1. Feuilles complètement glabres sur les deux faces (parfois avec un indument très petit sur la surface inférieure, visibles seulement sous microscope, ou de part et d’autre de la nervure médiane) ........................................................................................................ 2

1a. Feuilles avec au moins quelques trichomes évidents, parfois restreints à la nervure médiane ....................................................................................................................20

2. Feuilles généralement verticellées (trois par noeud, parfois une feuille manquante mais laissant toujours une cicatrice); forêts littorales orientales (parfois jusqu’à 200 m d’altitude).................................................................................................. 23. A. nitida View in CoL

2a. Feuilles alternes ou opposées à subopposées ...............................................................3

3. Limbe foliaire le plus grand mesurant au moins 15 cm de long ................................... 4

3a. Limbe foliaire le plus grand de moins de 15 cm long (rarement jusqu’à 17 cm chez A. oblongifolia View in CoL et A. littoralis View in CoL ).............................................................................................6

4. Pétiole robuste, d’env. 4-5 mm de diam.; jeunes rameaux avec une écorce liégeuse; forêts littorales et de basse altitude, de Maroantsetra au nord à Tampolo au sud ..................... ...............................................................................................................15. A. latifolia View in CoL

4a. Pétiole grêle, ne dépassant pas 2(-2.5) mm de diam.; jeunes rameaux sans écorce liégeuse; forêts humides .............................................................................................. 5

5. Feuille la plus grande avec env. 50-60 paires de nervures secondaires; fruit ellipsoïde, couvert d’un indument dense et court; Betampona........................ 2. A. betamponensis View in CoL

5a. Feuille la plus grande avec env. 30-45 pairs de nervures secondaires; fruit obovoïde, glabre; Lokobe et Tsaratanana.............................................................19. A. lokobensis View in CoL

6 (3). Feuilles elliptiques à ovées (rarement quelques unes légèrement obovales), plus larges dans la partie proximale .............................................................................................. 7

6a. Feuilles obovales à oblancéolées, presque toutes plus larges dans la partie distale .....11

7. Feuilles étroitement elliptiques à étroitement ovées (rarement quelques unes obovales), au moins (2.5-)3 fois plus longues que larges; forêts sèches, de l’Isalo à Andohahela..... ......................................................................................................... 27. A. phillipsonii View in CoL

7a. Feuilles elliptiques à ovées, parfois largement (rarement quelques unes légèrement obovales), pas plus de 1.5-2 fois plus longues que larges............................................. 8

24 Boissiera 71

8 (7). Fruits couverts d’un indument court et dense; fleurs 4-mères (rarement 5-mères); forêts humides, Marojejy et Anjanaharibe-dans le nord jusqu’à Ranomafana et Ivohibe dans le sud ................................................................................................... 17. A. lenticellata View in CoL

8a. Fruits glabres; fleurs 5-mères (inconnues chez A. darainensis View in CoL ) ..................................... 9

9. Feuilles orbiculaires à largement elliptiques (rarement quelques unes largement ovées), limbes pas plus de 2 fois plus longs que larges, subcoriaces, apex arrondi (rarement aigu); forêts sèches sur substrats calcaires, Ankarana, Bemaraha ................................. ........................................................................................................... 12. A. humbertii View in CoL

9a. Feuilles elliptiques à ovées (parfois quelques une largement ovées ou obovales), la plupart des limbes au moins 2 fois plus longs que larges, chartacées, apex aigu à acuminé ....................................................................................................................10

10. Limbe de la feuille la plus grande mesurant env. 6.5 cm de long, souvent discolore, la surface supérieure vert foncé à brun-chocolat, apex émarginé ou arrondi à aigu; forêts sèches occidentales (Bemaraha, Ketsepy, Antsalova, Marovoay, Mahajanga) ................... ................................................................................................................. 6. A. deflexa View in CoL

10a. Limbe de la feuille la plus grande env. 8 cm de long, concolore, la surface supérieure gris-vert sur le sec, apex acuminé; forêts humides, Daraina ................ 5. A. darainensis View in CoL

11 (6). Feuille (la plus grande) avec 40-70 paires de nervures secondaires; forêt humide orientale .................................................................................................. 30. A. sericea View in CoL

11a. Feuille (la plus grande) avec 10-40 paires de nervure secondaires .............................12

12. Inflorescences exclusivement axillaires; feuilles alternes, forêts littorales orientales, de Tampolo au nord jusqu’à Mahabo au sud ..............................................18. A. littoralis View in CoL

12a. Inflorescences terminales mais quelquefois aussi axillaires; feuilles opposées à subopposées. .............................................................................................................13

13. Nervures tertiaires invisibles sur la surface inférieure.................................................14

13a. Nervures tertiaires visible sur la surface inférieure, du moins vers la marge du limbe....16

14. Nervures secondaires étroitement espacées, 6-9 paires par cm au niveau de la partie médiane de la plus grande feuille, limbe glauque sur la face supérieure, presque toujours fortement discolore sur le sec, souvent brun-chocolaté sur la surface supérieure (parfois caché par la glaucescence); fruit velu (parfois glabrescent); forêts sèches, partie centrale des hauts plateaux, Itremo, Ibity, Ambatofinandrahana .......................... 3. A. buxifolia View in CoL

14a. Nervures secondaires largement espacées, 3-5 paires par cm au niveau de la portion médiane de la feuille la plus grande, limbe ni glauque (sauf un peu chez A. suarezensis View in CoL ) ni fortement discolore sur le sec, divers tons de vert mais jamais brun-chocolaté sur la surface supérieure; fruit glabre ..................................................................................15

15 (14).Feuille (la plus grande) avec 15-30 paires de nervures secondaires, à marge distinctement révolutée; arbres de 15-20 m de haut; forêts littorales et sèches, Mt. d’Ambre, sud de Vohémar et Masoala .............................................................................. 28. A. revoluta View in CoL

15a. Feuille (la plus grande) avec 12-15 paires de nervures secondaires, à marge légèrement épaissie mais pas révolutée; arbres de 10(-12) m de haut; forêts sèches et côtières du Nord, Antsiranana, Vohémar et Daraina .......................................... 31. A. suarezensis View in CoL

16 (13).Feuilles à nervures craspédodromes, toutes les nervures secondaires atteignant la marge (rarement quelques unes n’atteignant pas la marge chez A. oblongifolia View in CoL ).....................17

16a. Feuilles à nervures semi-craspédodromes, au moins quelques nervures secondaires ramifiées et n’atteignant pas la marge ........................................................................19

17. Feuille (la plus grande) avec 40-70 paires de nervures secondaires, limbe le plus grand de 9-15 cm long ..................................................................................... 30. A. sericea View in CoL

17a. Feuille (la plus grande) avec 15- 33 paires de nervures secondaires, limbe le plus grand de 2.5-7.5(-8.5) cm long ...........................................................................................18

18. Limbe de la feuille la plus grande d’au moins (4.5-) 5 cm de long, coriace, rigide; fruits mûrs de 2-3 × 1.4-1.6 cm ...................................................................... 8. A. ditimena View in CoL

18a. Limbe de la feuille la plus grande ne dépassant pas 4-(4.5) cm de long, subcoriace, non rigide; fruits mûrs 1-1.5 × 0.5-0.8 cm ...............................................22. A. minutifolia View in CoL

19 (16).Limbe de la feuille la plus grande de 8-14(-17) cm de long, pétiole env. 2 mm de diam.; fruits glabres; forêts sèches à humides, Nossi Be, Rég. d’Antsiranana ............................. ........................................................................................................ 24. A. oblongifolia View in CoL

19a. Limbe de la feuille la plus grande de 3.5-6 cm long, pétiole env. 0.5-1 mm de diam.; fruits à indument dense et court; forêts littorales et humides de basse altitude, Rég. de Tolagnaro ..........................................................................................7. A. delphinensis View in CoL

20 (1). Limbe de la feuille la plus grande d’au moins 10 cm de long.....................................21

20a. Limbe de la feuille la plus grande ne dépassant pas 10 cm de long............................34

21. Feuilles les plus grandes avec au moins 40 paires de nervures secondaires ................22

21a. Feuilles les plus grandes avec moins de 40 paires de nervures secondaires ................26

22. Inflorescences exclusivement axillaires ......................................................................23

22a. Inflorescences terminales et parfois aussi axillaires ....................................................24

23. Nervation tertiaire évidente, apex de la feuille plan; fruits glabres; forêts humides, Marojejy, Anjanaharibe-Sud, Masoala, Rantabe, Ambalabe ....................10. A. elongata View in CoL

23a. Nervation tertiaire absente, apex de la feuille généralement plié sur les spécimens séchés; fruits jeunes pubescents, glabrescents; forêts humides, Ambilobe, Ampondrabe, Ambato, Kalabenono-Galoka, Ambanja, Andavakoera, Tsaratanana............... 29. A. sambiranensis View in CoL

26 Boissiera 71

24 (22).Nervures secondaires de la partie médiane du limbe étroitement espacées, env. 5-6 par cm; fruit à indument dense; forêts humides orientales.............................. 30 A. sericea View in CoL

24a. Nervures secondaires de la partie médiane du limbe largement espacées, env. 3 par cm; fruit glabre ou à indument clairement visible.............................................................25

25. Apex de la feuille aigu à courtement acuminé (rarement quelques feuilles à apex arrondi); fruits à indument dense et court; fleurs 4-mères (rarement 5-mères); forêts humides.... ......................................................................................................... 17. A. lenticellata View in CoL

25a. Apex de la feuille obtus à arrondi ou rétus; fruits glabres; fleurs 5-mères; forêts sèches ......................................................................................................... 11. A. grandidieri View in CoL

26 (21).Inflorescences exclusivement axillaires ......................................................................27

26a. Inflorescences terminales et parfois aussi axillaires ....................................................29

27. Surface inférieure des feuilles densément laineuse, à indument brun clair, érigé à étalé ................................................................................................................ 34. A viguieri View in CoL

27a. Surface inférieure des feuilles glabres ou à indument court, épars .............................28

28. Pétiole le plus long mesurant 5-10 mm; forêts humides orientales de basse et moyenne altitude ................................................................................................. 16. A. lecomtei View in CoL

28a. Pétiole le plus long mesurant (18-) 20-30 mm; forêts littorales orientales, de Tampolo au nord jusqu’à Mahabo au sud..................................................................18. A. littoralis View in CoL

39 (26).Apex de la feuille longuement acuminé, presque caudé, limbe chartacé ....................30

29a. Apex de la feuille aigu à rétus ou émarginé, parfois courtement acuminé (acumen <1 cm de long), limbe chartacé à subcoriace ou coriace .......................................................32

30. Surface inférieure des feuilles densément laineuse à indument brun rougeâtre, nervures craspédodromes; forêts littorales orientales sur sable ou forêt de piémont sur latérite, Maroantsetra, Toamasina, Zahamena PN, Fort-Carnot ............................ 34. A. viguieri View in CoL

30a. Surface inférieure des feuilles glabres ou à indument blanc soyeux et épars, nervures semi-craspédodromes................................................................................................31

31. Surface supérieure des feuilles toujours glabres, limbe le plus large de 3.5(-4) cm de largeur; fruit velu; forêts humides orientales......................................17. A. lenticellata View in CoL

31a Surface supérieure des feuilles glabres ou à indument blanc apprimé, limbe le plus large d’au moins de 4 cm de largeur; fruit glabre; forêts humides du nord-est, Masoala, Maroantsetra ....................................................................................1. A. antongilensis View in CoL

32 (29).Feuilles à 10-20 paires de nervures secondaires largement espacées, seulement 1 ou 2 par cm sur la partie médiane du limbe; forêts humides de l’est et du nord-est Maroantsetra, Manongarivo, Andapa, Sambava, Ranomafana, Mantadia, Zahamena ...... ......................................................................................................... 32. A. thouvenotii View in CoL

32a. Feuilles à 20-40 paires de nervures étroitement espacées, 2-6 par cm au niveau de la partie médiane du limbe ...........................................................................................33

33. Feuilles obovales, jamais plus de 2.5 fois plus longues que larges, coriaces; inflorescences terminales et parfois aussi axillaires; fruits à indument dense; forêts humides orientales ............................................................................................................... 8. A. ditimena View in CoL

33a. Feuilles oblancéolées (parfois étroitement elliptiques), au moins 3 fois plus longues que larges, chartacées à coriaces, flexibles; inflorescences terminales et axillaires; fruits glabres; le long des cours d’eau, ouest et sud de Madagascar, Ambanja, Befandriana Nord, Ambalavao, Ihosy, Sakaraha, Amboasary, Ampanihy............... 11. A. grandidieri View in CoL

34 (20).Feuilles les plus grandes avec au moins 40 paires de nervures secondaires ................35

34a. Feuilles les plus grandes avec moins de 40 paires de nervures secondaires ................36

35. Nervures secondaires au niveau de la partie médiane du limbe étroitement espacées, env. 5-6 par cm; fruit à indument dense; forêts humides orientales ......... 30. A. sericea View in CoL

35a. Nervures secondaires au niveau de la partie médiane du limbe largement espacées, env. 3 par cm; fruit glabre; le long des cors d’eau, ouest et sud de Madagascar, Ambanja, Befandriana Nord, Ambalavao, Ihosy, Sakaraha, Amboasary, Ampanihy ........................ ......................................................................................................... 11. A. grandidieri View in CoL

36 (34).Feuilles les plus grandes avec un maximum de 20 paires de nervures secondaires.....37

36a. Feuilles les plus grandes avec plus de 20 paires de nervures secondaires ...................46

37. Feuilles elliptiques à orbiculaires ou légèrement ovées, la plus grande largeur au niveau de la partie médiane ou en-deça ................................................................................38

378a. Feuilles obovales à oblancéolées, la plus grande largeur au-delà de la partie médiane...40

38. Feuilles orbiculaires à largement elliptiques (rarement quelques feuilles largement ovées), pas plus de 2 fois plus longues que larges; forêt sèche, Ankarana, Bemaraha ..... ........................................................................................................... 12. A. humbertii View in CoL

38a. Feuilles elliptiques a étroitement elliptiques, au moins 2 fois plus longues que larges...39

39. Feuilles les plus grandes de 2.5-3.6 cm de large; forêts humides de transition, Manambato, Daraina, vallée de l’Androranga près de Sambava .................................. 4. A. capuronii View in CoL

39a. Feuilles les plus grandes de 1.5-2(-2.2) cm de large; forêts sèches, de l’Isalo à Andohahela .......................................................................................27. A. phillipsonii View in CoL

40 (37).Inflorescences exclusivement axillaires, inflorescences femelles avec un maximum de 5 fleurs (inflorescences males inconnues); forêts humides, Nosy Be, Ampasindava, Ambongo-Boina ................................................................................. 26. A. pauciflora View in CoL

40a. Inflorescences terminales et parfois aussi axillaires, fleurs 8-nombreuses (rarement seulement 5 fleurs dans les inflorescences femelles de A. suarezensis View in CoL ).........................41

28 Boissiera 71

41 (40).Limbe des feuilles les plus grandes 2-3(-4) cm de long; fruits 1-1.5 cm de long ............ ......................................................................................................... 22. A. minutifolia View in CoL

41a. Limbe des feuilles les plus grandes (3.5-) 4 cm de long; fruits 1.5-2.5 cm de long .....42

42. Fruits densément couverts d’un indument court; nervures secondaires semicraspédodrome, au moins quelques nervures ramifiées et n’atteignant pas la marge; forêts littorales, Rég. de Tolagnaro .....................................................7. A. delphinensis View in CoL

42a. Fruit glabres, nervures secondaires craspédodromes, toutes atteignent clairement la marge ........................................................................................................................43

43. Pétiole robuste, (1.5-) 2-3 mm de diam. sur les feuilles les plus grandes, nervures tertiaries distinctivement surélevées sur la surface inférieure; forêts humides de l’est et du nord-ouest, Ambanja, Marojejy, Anajanaharibe Sud, Ranomafana, Analamazaotra, Zahamena ........................................................................................ 32. A. thouvenotii View in CoL

43a. Pétiole grêle, pas plus de 1.5 mm de diam. sur les feuilles les plus grandes, nervures tertiaries indistinctes à faiblement surélevées sur la surface inférieure .......................44

44. Nervures secondaires étroitement espacées, 5-6 paires par cm au niveau de la partie médiane de la feuille la plus grande; forêts humides orientales, Analamazaotra, Ambalabe ............................................................................................... 20. A. louvelii View in CoL

44a. Nervures secondaires largement espacées, 3-4 paires par cm au niveau de la partie médiane de la feuille la plus grande ..........................................................................45

45. Inflorescences de 1-3(-3.5) cm de long; feuille à marge ondulée, nervures secondaires env. 10-12(-14) paires; forêts sèches et côtières du Nord, Antsiranana, Vohémar, Daraina ............................................................................................ 31. A. suarezensis View in CoL

45a. Inflorescences de 4-9 cm de long; feuille à marge foliaire plane ou faiblement ondulée, nervures secondaires 14 paires ou plus; forêts humides, Ankazobe, E de Fianarantsoa, Ranomafana, Andohahela (Parcelle n° 1), Ivohibe ..................................... 33. A. turkii View in CoL

46 (36).Nervures secondaires semi-craspédodromes, au moins quelques nervures ramifiées et n’atteignant pas la marge; fruit à indument visible.............................17. A. lenticellata View in CoL

46a. Nervures secondaires craspédodromes, atteignent toutes la marge; fruit glabre ou à indument évident ......................................................................................................47

47. Pétiole des feuilles les plus grandes d’au moins de 10 mm long.................................48

47a. Pétiole des feuilles les plus grandes de moins de 10 mm long....................................53

48. Inflorescences exclusivement axillaires; forêts littorales orientales, de Tampolo au nord jusqu’à Mahabo au sud..........................................................................18. A. littoralis View in CoL

48a. Inflorescences terminales et parfois aussi axillaires ....................................................49

49 (48).Nervures tertiaires à peine visibles, presque absentes; fleurs femelles à corolle glabre sur la surface externe.......................................................................... 20. A. longipetiolata View in CoL

49a. Nervures tertiaires visibles au moins sur une surface; corolle de la fleur femelle avec un indument sur la surface externe.................................................................................50

50. Feuilles pas plus de 2.5 fois plus longues que larges; fruits avec un indument évident ..................................................................................................................................51

50a. Feuilles au moins 3 fois plus longues que larges; fruits glabres ..................................52

51. Feuilles chartacées; forêts littorales, aux environs de Tolagnaro .........7. A. delphinensis View in CoL

51a. Feuilles coriaces; forêts humides orientales............................................. 8. A. ditimena View in CoL

52 (50).Feuilles obovales à elliptiques; à18-33paires de nervures secondaires, espacées de 1-4mm; forêts humides et sub-humides de transition du nord-est (Daraina, Vohémar)............... .............................................................................................................. 4. A. capuronii View in CoL

52a. Feuilles oblancéolées;de35-40-(47)paires de nervures secondaires,espacées de 0.5-5mm; forêts sèches et sub-humides de l’ouest et su sud-ouest .................... 11. A. grandidieri View in CoL

53 (47).Nervures secondaires faiblement saillantes à la face supérieure des feuilles; fruits glabres ......................................................................................................................54

53a. Nervures secondaires distinctement saillantes à la face supérieure des feuilles; fruits à indument évident ......................................................................................................60

54. Fruits de 3-3.5 cm de long, ellipsoïdes; axes des infructescence robustes, de 1.5-2 mm de diam.; feuilles les plus grandes mesurant 8-9 × 2.5 cm, apex aigu; forêts littorales, aux environs de Tolagnaro ........................................................................ 9. A. ellipticarpa View in CoL

54a. Fruits de 1.5-2.6 cm de long, ovoïdes ou ellipsoïdes; axes des infructescence grêles, env. 1(-1.2) mm de diam.; feuilles les plus grandes soit de moins de 8 × 2 cm, soit avec un apex arrondi à émarginé.......................................................................................55

55. Feuilles obovales à oblancéolées ................................................................................56

55a. Feuilles elliptiques à oblongues, rarement quelques-unes légèrement ovées ou obovales ....................................................................................................................59

56. Feuilles les plus grandes avec au moins 35 paires de nervures secondaires; forêts sèches .............................................................................................. 11. A. grandidieri View in CoL

56a. Feuilles les plus grandes avec pas plus de 25(-28) paires de nervures secondaires; forêts humides ....................................................................................................................57

57. Nervures secondaires largement espacées, 3-4 paires par cm au niveau de la partie médiane du limbe le plus grand; forêts humides, Ankazobe, à l’Est de Fianarantsoa, Ranomafana, Andohahela (Parcelle n° 1), Ivohibe ......................................... 33. A. turkii View in CoL

57a. Nervures secondaires étroitement espacées, 5-6 paires par cm au niveau de la partie médiane du limbe le plus grand ................................................................................58

30 Boissiera 71

58 (57).Limbe foliaire de la feuille la plus grande de 4(-4.2) cm de long; forêts humides, Analamazaotra, Ambalabe ...................................................................... 21. A. louvelii View in CoL

58a. Limbe foliaire de la feuille la plus grande de (4.8-) 5 cm de long; forêts semidécidues et h humides, Daraina ..................................................................................25. A. patrickii View in CoL

59 (55).Pétiole grêle, de moins de 1 mm de diam.; cotylédons fusionnés; arbres d’env. 3-12 m; forêts sèches, Isalo to Andohahela .................................................... 27. A. phillipsonii View in CoL

59a. Pétiole robuste, de 1.2-1.5 mm de diam.; cotylédons séparables; arbuste ou petits arbres de moins de 2 m de haut; forêts sclérophylles, Ibity ............................ 13. A. ibityensis View in CoL

60 (53).Feuilles glauques à la face supérieure, presque toujours fortement discolores sur le sec, souvent brun-chocolaté sur la face supérieure (parfois caché par la glaucescence); forêts sèches, partie centrale du haut plateau, Itremo, Ibity, Ambatofinandrahana................... ............................................................................................................... 3. A. buxifolia View in CoL

60a. Feuilles non glauques à la face supérieure, concolores ou faiblement discolores ........61

61. Feuilles pubérulentes à la surface supérieure; Rég. d’Itremo ............. 14. A. itremoensis View in CoL

61a. Feuilles glabres à la face supérieure ...........................................................................62

62. Feuilles chartacées; forêts littorales, aux environs de Tolagnaro .........7. A. delphinensis View in CoL

62a. Feuilles coriaces; forêts humides orientales............................................. 8. A. ditimena View in CoL

Table 1. MorphoLogicaL features distinguishing Abrahamia Randrian.& Lowry from Protorhus EngL.

  Abrahamia Protorhus
Fruit shape ELLipsoidaL or ovoid, radiaLLy symmetricaL Ovoid, mango-shaped or transverseLy obLong, asymmetricaL
Seed Resiniferous canaLs present, making the seed ruminate; cotyLedons inseparabLe Resiniferous canaLs Lacking, seed not ruminate; cotyLedons separabLe
Number of styles 1 (apicaLLy trifid or with 3 capitate stigmas) 3, sLightLy connate at the base or compLeteLy free
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