Combretaceae

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Combretaceae

Content

<<<Flowers>Ovary>Ovules>Ovule number

(1-)2-71
1. 005-007-002-001

<<<Flowers>Ovary>Ovules>Position

pendulous2
2. 005-007-002-002

<<<Flowers>Ovary>Style>Style number

13
3. 005-007-003-001

<<<Flowers>Stamens>Anthers>Locule number

4-locular4
4. 005-005-001-001

<<<Fruits>Seeds>Cotyledons>Cotyledon number

usually 25
5. 006-003-003-001

<<Flowers>Hypanthium>Prophyllum number

26
6. 005-002-001

<<Flowers>Ovary>Locule number

1-locular7
7. 005-007-001

<<Flowers>Ovary>Ovules

ovules , apical, ,8
8. 005-007-002

<<Flowers>Ovary>Style

style , with usually punctiform stigma.9
9. 005-007-003

<<Flowers>Petals>Petal number

4-5, or often 010
10. 005-004-001

<<Flowers>Sepals>Sepal number

4-511
11. 005-003-001

<<Flowers>Stamens>Anthers

anthers , dorsifixed, usually versatile, sometimes adnate;12
12. 005-005-001

<<Fruits>Seeds>Cotyledons

cotyledons , dorsiventral and variously folded or twisted (mostly spirally convolute or irregularly complicate), rarely conduplicate.13
13. 006-003-003

<<Fruits>Seeds>Endosperm

endosperm absent,14
14. 006-003-002

<<Fruits>Seeds>Seed number

115
15. 006-003-001

<Flowers>Hypanthium

hypanthium (receptacle) surrounding ovary (lower hypanthium) and extended beyond into saucer- to tube-shaped upper hypanthium bearing stamens and perianth, with prophylls (bracteoles) fused to lower hypanthium in Laguncularieae;16
16. 005-002

<Flowers>Merosity

4- to 5-merous17
17. 005-001

<Flowers>Nectary disc

nectariferous disk, continuous or as separate lobes, often present at base of or at some distance up upper hypanthium;18
18. 005-006

<Flowers>Ovary

ovary ,19
19. 005-007

<Flowers>Petals

petals , free, alternating with sepals, usually borne at or near tip of upper hypanthium, often small, sometimes conspicuous;20
20. 005-004

<Flowers>Sepals

sepals , borne at tip of upper hypanthium, sometimes vestigial, rarely accrescent;21
21. 005-003

<Flowers>Stamens

stamens usually twice as many as sepals, borne inside upper hypanthium usually at two levels, sometimes as many as sepals, antipetalous [or antisepalous] whorl missing, exserted or included,22
22. 005-005

<Fruits>Seeds

seed ,23
23. 006-003

<Fruits>Texture

papery to leathery24
24. 006-002

<Fruits>Wing number

2-525
25. 006-001

<Indumentum>Glands

short, capitate stalked glands, and subsessile peltate 'scales'.26
26. 003-002

<Indumentum>Hairs

pointed hairs with a distinctive basal compartment (‘combretaceous hairs’) alone or with glandular hairs of one of two types27
27. 003-001

<Inflorescences>Bracts

bracts simple, usually caducuous.28
28. 004-002

<Inflorescences>Position

axillary or terminal29
29. 004-001

Description

Trees, shrubs or lianes, sometimes mangroves, not spiny. Leaves opposite (or whorled) or spiral (often ‘alternate’), petiolate, simple, entire, without stipules, often with a pair of petiolar glands. Indumentum almost always of unicellular, slender, thick-walled, pointed hairs with a distinctive basal compartment (‘combretaceous hairs’) alone or with glandular hairs of one of two types: short, capitate stalked glands, and subsessile peltate 'scales'. Inflorescences axillary or terminal, capitate to expanded, of simple or paniculate spikes or less often racemes; bracts simple, usually caducuous. Flowers bisexual or bisexual and male in same inflorescence or sometimes dioecious, 4- to 5-merous, actinomorphic or sometimes weakly zygomorphic, epigynous; hypanthium (receptacle) surrounding ovary (lower hypanthium) and extended beyond into saucer- to tube-shaped upper hypanthium bearing stamens and perianth, with 2 prophylls (bracteoles) fused to lower hypanthium in Laguncularieae; sepals 4-5, borne at tip of upper hypanthium, sometimes vestigial, rarely accrescent; petals 4-5, or often 0, free, alternating with sepals, usually borne at or near tip of upper hypanthium, often small, sometimes conspicuous; stamens usually twice as many as sepals, borne inside upper hypanthium usually at two levels, sometimes as many as sepals, antipetalous [or antisepalous] whorl missing, exserted or included, anthers 4-locular, dorsifixed, usually versatile, sometimes adnate; nectariferous disk, continuous or as separate lobes, often present at base of or at some distance up upper hypanthium; ovary 1-locular, ovules (1-)2-7, apical, pendulous, style 1, with usually punctiform stigma. Fruits indehiscent, with dry or spongy to succulent wall, often with 2-5papery to leathery wings; seed 1, endosperm absent, cotyledons usually 2, dorsiventral and variously folded or twisted (mostly spirally convolute or irregularly complicate), rarely conduplicate.

Discussion

Unless otherwise stated, leaf length excludes the petiole, and flower length includes the ovary and calyx lobes but excludes the petals, stamens and style, and upper hypanthium length also includes the calyx lobes; upper hypanthium width is taken at junction of hypanthium and calyx lobes in Combretum, but includes the calyx lobes in Terminalia. The lower hypanthium is taken to include the ovary, any pedicel-like proximal region present, and the whole of any solid distal extension below the expanded upper hypanthium. All flower measurements are taken from boiled material; dried material is obviously smaller. Fruit length includes pseudostipe and beak where present, taken at midline; lateral extent of wing is described as wing-width. Leaf venation terminology follows Hickey (1973); descriptions refer to leaf lower surface unless indicated otherwise.

Distribution

Americas present, Guianas present, New World present, Southern America: Venezuela (Venezuela present), throughout the tropics and subtropics of the whole world present
About 500 species in 14 genera, distributed throughout the tropics and subtropics of the whole world; 5 genera occur in the New World and in the Guianas; of the 85 species in the Americas, 31 occur in the Guianas (plus 1 nearby in Venezuela).

FAMILY DESCRIPTION

Growth rings absent to distinct.
Vessels diffuse, round to oval, solitary and in short radial multiples of 2-3(-10); typical vessels in Combretum rarely intermingled with very narrow vessels and vascular tracheids in small clusters; mean tangential diameter 87-170 μm; 3-25 per mm². Perforations simple with horizontal to oblique end walls. Intervessel pits vestured, alternate, round to polygonal, (5-)6-9(-11) μm, rarely coalescent; vessel-ray pits similar to intervessel pits but half-bordered, sometimes in horizontal arrangement, elongate, rarely tending to scalariform or coalescent.
Rays exclusively uniseriate in Laguncularia, nearly so in Buchenavia, Combretum, Conocarpus and Terminalia. Rays composed of square and upright, and weakly procumbent cells in Buchenavia and Laguncularia.In Conocarpus upright to procumbent cells are intermingled. In Combretum and Terminalia, cells are procumbent (with infrequent square and upright cells and upright marginal cells); 7-11 per mm, but up to 20 in Combretum and Terminalia. Perforated ray cells present in Combretum.
Parenchyma apotracheal diffuse, sometimes in marginal bands or absent; paratracheal vasicentric to aliform-confluent, sometimes in bands of 2-5 cells wide; strands of (3-)4-7(-8) cells.
Ground tissue fibres thin-walled to very thick-walled; non-septate in Laguncularia, but septate in Buchenavia and Conocarpus. In Combretum and Terminalia, both septate and non-septate fibres can be found; pits simple to minutely bordered, mainly on radial cell walls.
Crystals present in rays and/or axial parenchyma, as small or large rhombic crystals, or as elongated rod- to styloid-like crystals, sometimes scanty (Table 1).
Table 1. Occurrence of crystals in Guianan COMBRETACEAE. Data between brackets: crystals are present in some species. ?: crystals have been reported by van Vliet (1979), but not confirmed in this study.raysaxial parenchyma
Buchenavia(rhombic, elongate)
Combretumsmall, rhombic in large ideoblastslarge, rhombic, in large ideoblasts
Conocarpuslarge, blunt prismatic, in radial rows
Laguncularialarge, prismatic to elongate
Terminalia(large, blunt prismatic)scanty rhombic; druses in T.catappa?

Flowers

Flowers bisexual or bisexual and male in same inflorescence or sometimes dioecious, , actinomorphic or sometimes weakly zygomorphic, epigynous;30
30. 005

Fruits

Fruits indehiscent, with dry or spongy to succulent wall, often with wings;31
31. 006

Habit

Trees, shrubs or lianes, sometimes mangroves, not spiny.32
32. 001

INTRODUCTION

The most recent global treatment of the wood anatomy of the COMBRETACEAE is by van Vliet (1979). Therefore, his descriptions were used as a basis for this chapter, supplemented by our own observations.
In van Vliet’s thorough study, each generic description was followed by an extensive discussion of the individual characters, their taxonomical value and their correlation with habit and habitat, and the classification of the family. Most of those data and conclusions go beyond the scope of this Flora. For example: when studying the bordered intervessel pits in COMBRETACEAE and allied families, van Vliet (1978, 1979) was able to distinguish between 2 main types, based on size of the vestures and their position in the pit chamber. One intervessel pit type (type A) was characteristic for the genus Strephonema Hook.f. (not studied here). The other type (type B), found in all other genera of COMBRETACEAE, was subdivided into 3 forms, but these subtypes were not correlated with other woodanatomical characters, nor did van Vliet find any correlation with the taxonomy of the family. Only in Terminalia the full variation of type B was found. As the use of a scanning electron microscope is essential to decide on the type of vesturing in a given sample, we refrained from describing the vesture types here. Moreover, a recapitulation here of van Vliet’s discussion of anatomical variation in relation to habit and habitat, but restricted to Guianan material only, would not contribute to, let alone improve, his conclusions and thus the reader is referred to the original work (van Vliet 1978).
Other quantitative characters that may be difficult to measure accurately from sections, such as vessel member length, ray height, are not given here, notwithstanding the fact they may have diagnostic value.
Van Vliet reports 2 distinct types of vessel elements for Combretum. Besides the elements typical for COMBRETACEAE, very narrow elements, often associated with vascular tracheids are present. In the Guianan species of Combretum, however, they are rarely found.
Descriptive terms and measurements are in accordance to the IAWA-list of microscopic features (1989).

Individuals Association

Guyana, Hoffman, B., et al. 1000 Guyana, Clarke, D., et al. 6437 Guyana, Watson, A.G. 32 Guyana, Mutchnick, P., et al. 1269 Weitzman, A.L., et al. 247 Poiteau, P.A. s.n. Suriname, Hostmann, F.W. 744 Guyana, Görts-van Rijn, A.R.A., et al. 294 Guyana, Gillespie, L.J., et al. 2409 Suriname, BW 1488 Suriname, Lanjouw, J. & J.C. Lindeman 1037 Mori, S.A., et al. 15121 Oldeman, R.A.A. B-1531 Richard, L.C.M. s.n. Bordenave, B. 850 Guyana, Clarke, D., et al. 3926 Suriname, Hostmann, F.W. & A. Kappler 138 Guyana, Pipoly, J.J., et al. 7459 Prévost, M.F., et al. 3564 Guyana, Clarke, D., et al. 8009 Oldeman, R.A.A. B-2157 Suriname, Wullschlägel, H.R. 152 Oldeman, R.A.A. B-2308 Guyana, Clarke, D., et al. 2232 Guyana, Anderson, C.W. 390 Sabatier, D., et al. 3515 Guyana, Hoffman, B., et al. 2753 Guyana, Henkel, T.W., et al. 369 Suriname, Irwin, H.S., et al. 55359 Martin, J. 197 Martin, J. s.n. Guyana, Forest Dept. British Guiana 7779 Sabatier, D., et al. 2299 Guyana, Forest Dept. British Guiana 4329 Granville, J.J. de, et al. B-5267 Guyana, Forest Dept. British Guiana 4006 Guyana, Polak, M., et al. 119 Guyana, Forest Dept. British Guiana 3581 p.p. Guyana, Forest Dept. British Guiana 7231 Guyana, Cruz, J.S. de la 2680 Guyana, Hohenkerk, L.S. 136B Suriname, Schulz, J.P. 7628 Guyana, Mutchnick, P., et al. 1015 Guyana, Jenman, G.S. 4347 Guyana, Persaud, A.C. 223 Guyana, Pipoly, J.J., et al. 7438 Guyana, Meyer, P.E. 45 Guyana, Hitchcock, A.S. 17166 Suriname, Lanjouw, J. & J.C. Lindeman 1059 Guyana, Jenman, G.S. 5076 Guyana, Persaud, A.C. 389 Barrabé, L., et al. 174 Sabatier, D., et al. 1151 Mori, S.A., et al. 15343 Guyana, Persaud, A.C. 56 Guyana, Cruz, J.S. de la 1094 Suriname, Stahel, G. 363 Guyana, Sandwith, N.Y. 619 Raynal-Roques, A., et al. 24781 Guyana, McDowell, T., et al. 2143 Suriname, Kramer, K.U., et al. 2761 Guyana, Hoffman, B., et al. 3419 Oldeman, R.A.A. B-3682 Oldeman, R.A.A. 2825 Suriname, Tulleken, J.E. 409 Suriname, Mennega, A.M.W. 532 Guyana, Cruz, J.S. de la 4580 Guyana, Andel, T. van, et al. 1280 Suriname, Splitgerber, F.L. 112 Suriname, Hekking, W.H.A. 1077 Service Forestier 7913 Guyana, Forest Dept. British Guiana 6162 Leblond, J.B. 437 Suriname, Gonggrijp, J.W. 3689 Boom, B.M., et al. 2237 Leblond, J.B. s.n. Sabatier, D., et al. 2179 Guyana, Pipoly, J.J., et al. 7795 Suriname, Maguire, B. 23623 Suriname, Maguire, B. 23809 Guyana, Forest Dept. British Guiana 4811 Guyana, Henkel, T.W., et al. 2126 Aublet, J.B.C.F. s.n. Guyana, McDowell, T., et al. 2846 Suriname, Sastre, C., et al. 8126 Oldeman, R.A.A. 282 Guyana, Stergios, B., et al. 5272 Service Forestier 4047 Guyana, Gillespie, L.J., et al. 1380 Mori, S.A., et al. 8948 Guyana, Clarke, D., et al. 4771 Guyana, Clarke, D., et al. 4756 Guyana, Hoffman, B., et al. 4769 Sagot, P.A. 241 Guyana, Clarke, D., et al. 2230 Guyana, Maguire, B., et al. 32640 Granville, J.J. de, et al. 6852 Suriname, Heyde, N.M., et al. 422 Guyana, Forest Dept. British Guiana 5418 Guyana, McDowell, T., et al. 2302 Guyana, Mutchnick, P., et al. 770 Suriname, Heyde, N.M., et al. 404 Guyana, Gleason, H.A. 397 Guyana, Henkel, T.W., et al. 5134 Cremers, G., et al. 8401 Aubréville, A. 333 Guyana, Graham, J. 355 Guyana, Gillespie, L.J., et al. 1447 Garnier, F.A. 195 Guyana, Sandwith, N.Y. 104 Guyana, Clarke, D., et al. 3924 Patris, J.B. s.n. Suriname, BW 1312 Guyana, Pipoly, J.J., et al. 11777 Grenand, P. 1734 Poiteau, P.A. s.n. Hallé, F. 539 Grenand, P. 3014 Guyana, Clarke, D., et al. 271 Guyana, Gillespie, L.J., et al. 1386 Mélinon, E.M. 45 Sabatier, D., et al. 3355 Richard, L.C.M. s.n. Guyana, Cruz, J.S. de la 1509 Suriname, Stahel, G. 93 Sagot, P.A. s.n. Guyana, Dirven, J.G.P. LP194 Oldeman, R.A.A. B-1149 Guyana, Gillespie, L.J., et al. 1143 Suriname, Donselaar, J. van 1522 Suriname, Lems, K. 5100 Lemoine, S.V. 7897 Billiet, F., et al. 1897 Suriname, Lindeman, J.C. 9820 Fleury, M. 1121 Granville, J.J. de, et al. 8176 Oldeman, R.A.A. 2451 Sabatier, D., et al. 2298 Guyana, Clarke, D., et al. 7054 Oldeman, R.A.A. 2473 Guyana, Clarke, D., et al. 2757 Guyana, Mori, S.A., et al. 8153 Guyana, Forest Dept. British Guiana 7538 Aublet, J.B.C.F. 281 Oldeman, R.A.A. 2252 Guyana, Cruz, J.S. de la 2365 Prévost, M.F., et al. 977 Guyana, Pipoly, J.J., et al. 11717 Raynal-Roques, A., et al. 24780 Guyana, Persaud, A.C. 187 Guyana, Forest Dept. British Guiana 5915 Guyana, Clarke, D., et al. 7731 Guyana, Forest Dept. British Guiana 2252 Guyana, Mutchnick, P., et al. 988 Suriname, Went, F.A.F.C. 126 Mori, S.A., et al. 20763 Guyana, Mutchnick, P., et al. 373 Suriname, Went, F.A.F.C. 484 Guyana, Anderson, C.W. s.n. Sabatier, D., et al. 2283 Suriname, Irwin, H.S., et al. 55559 Guyana, Jansen-Jacobs, M.J., et al. 3407 Guyana, Jenman, G.S. 7240 Sagot, P.A. 795 Barrier, S. 5059 Granville, J.J. de, et al. 7153 Sabatier, D., et al. 4188 Guyana, Hoffman, B., et al. 576 Guyana, Forest Dept. British Guiana 4138 Oldeman, R.A.A. B-1421 Guyana, Henkel, T.W., et al. 2663 Guyana, Clarke, D., et al. 1378 Suriname, Lanjouw, J. & J.C. Lindeman 3460 Mori, S.A., et al. 20941 Guyana, Hahn, W.J., et al. 4807 Guyana, Reinders, M.A., et al. 159 Guyana, Smith, A.C. 3033 Suriname, Hugh-Jones, D.H. 50 Guyana, Gillespie, L.J., et al. 1369 Sabatier, D., et al. 1679 Oldeman, R.A.A. B-3611 Guyana, Jansen-Jacobs, M.J., et al. 2240 Prévost, M.F., et al. 3703 Guyana, Görts-van Rijn, A.R.A., et al. 482 Oldeman, R.A.A. 2981 Poiteau, P.A. s.n. Perrottet, G.S. s.n. Suriname, Maguire, B. 24067 Oldeman, R.A.A. 2516 Prévost, M.F., et al. 1312 Oldeman, R.A.A. B-3151 Guyana, Maguire, B., et al. 23355 Boom, B.M., et al. 2134 Guyana, Gillespie, L.J., et al. 1418 Guyana, Jansen-Jacobs, M.J., et al. 288 Suriname, Rombouts, H.E. 62 Sabatier, D., et al. 2627 Broadway, W.E. 3110 Guyana, Cruz, J.S. de la 3307 Fleury, M. 1154 Guyana, Archer, W.A. 2404 Guyana, Forest Dept. British Guiana 3775 Suriname, Mennega, A.M.W. 541 Lemoine, S.V. 7883 Guyana, Forest Dept. British Guiana 2705 Suriname, Versteeg, G.M. 540 Sabatier, D., et al. 2582 Guyana, Anderson, C.W. 749 Oldeman, R.A.A. 1323 Mori, S.A., et al. 23485 Guyana, Mutchnick, P., et al. 506 Suriname, Hostmann, F.W. 665 Suriname, Lanjouw, J. & J.C. Lindeman 1865 Richard, L.C.M. s.n. Guyana, Henkel, T.W., et al. 575 Suriname, Focke, H.C. 400 Suriname, Stahel, G. 363a Suriname, Hostmann, F.W. & A. Kappler 825 Guyana, McDowell, T., et al. 3329 Guyana, Leechman, A. s.n. Mélinon, E.M. 444 Guyana, Hoffman, B., et al. 732 Guyana, Henkel, T.W., et al. 1986 Suriname, Florschütz, P.A., et al. 2162 Guyana, Henkel, T.W., et al. 3353 Suriname, Jansma, R. LBB 15891 Guyana, Jansen-Jacobs, M.J., et al. 5706 Oldeman, R.A.A. 1435 Suriname, Irwin, H.S., et al. 55447 Guyana, Polak, M., et al. 521 Suriname, Wullschlägel, H.R. 1445 Sagot, P.A. 1 Suriname, Schulz, J.P. 7983 Suriname, Lanjouw, J. & J.C. Lindeman 610 Sagot, P.A. 242 Sabatier, D., et al. 1681 Guyana, Smith, A.C. 2833 Guyana, Tiwahri, S., et al. 961 Oldeman, R.A.A. B-2552 Suriname, Mori, S.A., et al. 8450 Richard, L.C.M. s.n. Granville, J.J. de, et al. B-4792 Guyana, Maguire, B., et al. 32640 Guyana, Maas, P.J.M., et al. 4059 Oldeman, R.A.A. T-324 Guyana, Forest Dept. British Guiana 4352 Guyana, Mutchnick, P., et al. 565 Lescure, J.P. 689 Guyana, Cruz, J.S. de la 1947 Guyana, Forest Dept. British Guiana 2378 Guyana, Jansen-Jacobs, M.J., et al. 2898 Guyana, Schomburgk, Ro. ser.I, 272 Prévost, M.F., et al. 4357 Guyana, Clarke, D., et al. 5074 Guyana, Henkel, T.W., et al. 3222 Guyana, Jansen-Jacobs, M.J., et al. 289 Guyana, Atkinson, D.J. 555 Guyana, Clarke, D., et al. 4778 Guyana, Henkel, T.W., et al. 564 Suriname, BBS 158 Mori, S.A., et al. 22642 Guyana, Cruz, J.S. de la 1488 Guyana, Hoffman, B., et al. 4566 Suriname, Samuels, J.A. 142 Suriname, Went, F.A.F.C. 123 Suriname, Hostmann, F.W. 1311 Guyana, Clarke, D., et al. 1380 Hallé, F. 2772 Suriname, Sauvain, M. 157 Lemée, A.M.V. s.n. Guyana, Jansen-Jacobs, M.J., et al. 4337 Hallé, F. 538 Benoist, R. 1256 Oldeman, R.A.A. 2652 Guyana, Cruz, J.S. de la 1333 Mélinon, E.M. s.n. Guyana, Tiwahri, S., et al. 376 Suriname, Jiménez-Saa, U. 1513 Suriname, Maas, P.J.M., et al. LBB 10834 Suriname, BW 2026 Leprieur, F.R.M. s.n. Guyana, Clarke, D., et al. 6497 Guyana, Grewal, M.S., et al. 335 Suriname, Focke, H.C. 829 Guyana, Cruz, J.S. de la 4258 Sabatier, D., et al. 575 Suriname, Kock, C. s.n. Guyana, Jenman, G.S. 1326 Suriname, Hostmann, F.W. & A. Kappler Suriname, Vreden, C., LBB 15774 Mori, S.A., et al. 25074 Guyana, Hoffman, B., et al. 1478 Suriname, Lindeman, J.C. & A.R.A. Görts et al. 631 Guyana, Tillett, S.S., et al. 45805 Guyana, Hoffman, B., et al. 1299 Oldeman, R.A.A. B-2136 Suriname, BW 5707 Guyana, Hoffman, B., et al. 2741 Guyana, Polak, M., et al. 302 Guyana, Forest Dept. British Guiana 5481 Prévost, M.F., et al. 329 Suriname, Mennega, A.M.W. 215 Leblond, J.B. s.n. Martin, J. s.n. Guyana, Forest Dept. British Guiana 5058 Suriname, Kegel, H.A.H. 979 Service Forestier 7897 Suriname, Lanjouw, J. 931 Suriname, Splitgerber, F.L. 354 Suriname, Heyde, N.M., et al. 74 Grenand, P. 1293 Suriname, Outer, R.W. den 969 Leprieur, F.R.M. s.n. Hoff, M. 5012 Guyana, Parker, C.S. s.n. Guyana, Clarke, D., et al. 3463 Suriname, Irwin, H.S., et al. 55789 Guyana, Hollowell, T.H., et al. 635 Guyana, Schomburgk, Ro. ser. I, 87 pro parte Loubry, D. 1949 Guyana, Mutchnick, P., et al. 1279 Sabatier, D., et al. 2264 Suriname, Heyde, N.M., et al. 24 Black, G.A., et al. 54-17593 Guyana, Henkel, T.W., et al. 3201 Guyana, Jenman, G.S. 4997 Oldeman, R.A.A. B-762 Oldeman, R.A.A. 1547 Guyana, Cruz, J.S. de la 1863 Guyana, Jenman, G.S. 3783 Suriname, Gonggrijp, J.W. 30 Guyana, Cruz, J.S. de la 3425 Guyana, Görts-van Rijn, A.R.A., et al. 491 Guyana, Cook, C.D.K. 247 Guyana, Hancock, J. s.n. Suriname, Rombouts, H.E. 701 Guyana, Forest Dept. British Guiana 7954 Suriname, Hekking, W.H.A. 797 Oldeman, R.A.A. B-1377 Grenand, P. 1303 Guyana, Watson, A.G. 1939 Oldeman, R.A.A. 1850 Guyana, Henkel, T.W., et al. 5129 Suriname, Lindeman, J.C. 3885 Desfontaines, R.L. s.n. Suriname, Elburg, J.P., LBB 9623 Suriname, BW 2035 Guyana, Jansen-Jacobs, M.J., et al. 146 Sagot, P.A. s.n. Oldeman, R.A.A. B-3403 Suriname, Sauvain, M. 735 Guyana, Maguire, B., et al. 23499 Suriname, Lindeman, J.C. 3812 Suriname, Kegel, H.A.H. 1191 Guyana, Clarke, D., et al. 3807 Sastre, C., et al. 8129 Broadway, W.E. 536 Suriname, Focke, H.C. 407 Mélinon, E.M. 206 Guyana, Maguire, B., et al. 23551 Sabatier, D., et al. 2309 Cremers, G., et al. 13732 Fleury, M. 356 Martin, J. s.n. Guyana, Mutchnick, P., et al. 458 Suriname, Mennega, A.M.W. 235 Guyana, Pennington, T.D., et al. 345 Guyana, Clarke, D., et al. 8712 Suriname, Teunissen, P.A. LBB 15484 Benoist, R. 2 Service Eaux et Forêts 4050 Sagot, P.A. 944 Guyana, Henkel, T.W., et al. 3657 Guyana, Gillespie, L.J., et al. 2194 Guyana, Mutchnick, P., et al. 512 Guyana, Jenman, G.S. 7455 Oldeman, R.A.A. B-2472 Leblond, J.B. 117 Mori, S.A., et al. 18625 Guyana, Jansen-Jacobs, M.J., et al. 3299 Suriname, Elburg, J.P., LBB 9820 Guyana, Stergios, B., et al. 5294 Aubréville, A. 222 Barrier, S. 5080 Guyana, Cruz, J.S. de la 2428 Guyana, Forest Dept. British Guiana 5065 Guyana, McDowell, T., et al. 1829 Suriname, Acevedo-Rodríguez, P., et al. 6160 Sabatier, D., et al. 1019 Guyana, Cruz, J.S. de la 2979 Suriname, Lanjouw, J. & J.C. Lindeman 1861 Guyana, Cruz, J.S. de la 3566 Hoff, M. 5105 Guyana, Forest Dept. British Guiana 4222 Suriname, Lanjouw, J. & J.C. Lindeman 1915 Suriname, Lanjouw, J. & J.C. Lindeman 1992 Guyana, Cook, C.D.K. 248 Suriname, Wessels Boer, J.G. 232 Guyana, Mutchnick, P., et al. 440 Guyana, Mutchnick, P., et al. 1295 Guyana, Hitchcock, A.S. 17516 Guyana, Granville, J.J. de, et al. 11702 Guyana, Clarke, D., et al. 3873 Guyana, Acevedo-Rodríguez, P., et al. 3302 Leprieur, F.R.M. s.n. Guyana, Jenman, G.S. 5043 Guyana, Jansen-Jacobs, M.J., et al. 462 Guyana, Chanderbali, A. 17 Guyana, Forest Dept. British Guiana 5096 Boom, B.M., et al. 2118 Guyana, Clarke, D., et al. 3787 Guyana, Jenman, G.S. 4359 Irwin, H.S., et al. 48328 Grenand, P. 78 Prévost, M.F., et al. 1781 Sabatier, D., et al. 4677 Guyana, Mutchnick, P., et al. 835 Geay, F. 1904 Sastre, C., et al. 8199 Guyana, Henkel, T.W., et al. 1978 Suriname, Hostmann, F.W. s.n. Bordenave, B. 826 Mori, S.A., et al. 22128 Oldeman, R.A.A. B-1400 Guyana, Schomburgk, Ro. ser. II, 872 Suriname, Geyskes, D.C. s.n. Bordenave, B. 1014 Guyana, Hoffman, B., et al. 1027 Mélinon, E.M. 291 Suriname, Versteeg, G.M. 553 Guyana, Gillespie, L.J., et al. 1665 Guyana, Hoffman, B., et al. 1210 Bois 7585 Guyana, Hoffman, B., et al. 2437 Mélinon, E.M. 69 Mori, S.A., et al. 15300 Mori, S.A., et al. 23905 Guyana, Forest Dept. British Guiana 2152 Bordenave, B. 855B Mélinon, E.M. 308 Guyana, Clarke, D., et al. 6661 Suriname, Kramer, K.U., et al. 2088 Sagot, P.A. 794 Suriname, Florschütz, P.A., et al. 1131 Guyana, Henkel, T.W., et al. 3399 Guyana, Cruz, J.S. de la 1440 Guyana, Stoffers, A.L., et al. 348 Prévost, M.F., et al. 2162 Hallé, F. 545 Sabatier, D., et al. 2837 Guyana, Cruz, J.S. de la 3713 Guyana, Davis, T.A.W. 1062 Oldeman, R.A.A. B-1783 Guyana, Schomburgk, Ro. ser. I, 87 pro parte Service Forestier 4172 Guyana, Parker, C.S. s.n. Feuillet, C. 2366 Mori, S.A., et al. 23905 Hallé, F. 600 Oldeman, R.A.A. B-2809 Foresta, H. de 476 Wachenheim, H. 52 Sabatier, D., et al. 3679 Guyana, Hoffman, B., et al. 1320 Suriname, Focke, H.C. 800 Suriname, Rolander, D. s.n. Grenand, P. 1401 Guyana, Persaud, A.C. 35 Sagot, P.A. 254 Oldeman, R.A.A. 2661 Guyana, Hohenkerk, L.S. 678 Bordenave, B. 784 Guyana, Henkel, T.W., et al. 2078 Guyana, Clarke, D., et al. 3531 Guyana, Lanjouw, J., et al. 874 Guyana, Hoffman, B., et al. 2743 Suriname, Versteeg, G.M. 707 Suriname, Geyskes, D.C. s.n. Guyana, Jansen-Jacobs, M.J., et al. 2878 Suriname, Donselaar, J. van 1510 Billiet, F., et al. 4496 Guyana, Pipoly, J.J., et al. 11751 Guyana, Jansen-Jacobs, M.J., et al. 5689 Guyana, Kelloff, C.L., et al. 1058 Guyana, Maguire, B., et al. 40656 Guyana, Cruz, J.S. de la 4104 Leprieur, F.R.M. s.n. Oldeman, R.A.A. 97 Service Forestier s.n. Guyana, Jansen-Jacobs, M.J., et al. 1573 Guyana, Jansen-Jacobs, M.J., et al. 1719 Guyana, Anderson, C.W. 79 Suriname, Hostmann, F.W. & A. Kappler 1151 Guyana, Archer, W.A. 2384 Guyana, Clarke, D., et al. 5060 Guyana, Henkel, T.W., et al. 3145 Mori, S.A., et al. 22140 Oldeman, R.A.A. B-2648 Suriname, BW 4661 Guyana, Martyn, E.B. 247 Guyana, Henkel, T.W., et al. 5270 Guyana, Maguire, B., et al. 23313 Guyana, Knapp, S., et al. 2780 Sagot, P.A. 255 Guyana, Maas, P.J.M., et al. 5570 Guyana, Gleason, H.A. 401 Granville, J.J. de, et al. 9600 Guyana, Jenman, G.S. 4890 Geay, F. 1903 Guyana, Gleason, H.A. 338 Service Forestier 7774 Guyana, Henkel, T.W., et al. 3124 Guyana, Jansen-Jacobs, M.J., et al. 2191 Granville, J.J. de, et al. 3948 Guyana, Omawale, et al. 42 Suriname, Kappler, A. s.n. Guyana, Jansen-Jacobs, M.J., et al. 5804 Guyana, Henkel, T.W., et al. 2434 Leblond, J.B. 403 Oldeman, R.A.A. B-673 Sagot, P.A. 1007 Guyana, Gillespie, L.J., et al. 2439 Guyana, Henkel, T.W., et al. 3463 Guyana, Forest Dept. British Guiana 3800 Suriname, Granville, J.J. de, et al. 11889 Suriname, Florschütz, P.A., et al. 148 Mori, S.A., et al. 15095 Guyana, Henkel, T.W., et al. 4570 Guyana, Polak, M., et al. 316 Guyana, Gillespie, L.J., et al. 1191 Guyana, Gillespie, L.J., et al. 2436 Billiet, F., et al. 1260 Petrov, I. 234 Gabriel, A. 18002 Hallé, F. 4013 Guyana, Appun, C.F. 288 Guyana, McDowell, T., et al. 2186 Guyana, Watson, A.G. 39 Suriname, BW 5471 Suriname, Schulz, J.P. 8701 Sabatier, D., et al. 2692 Guyana, Andel, T. van, et al. 3008 Moretti, C. 634 Guyana, Gillespie, L.J., et al. 1185 Suriname, BW 374 Aublet, J.B.C.F. s.n. Guyana, Jenman, G.S. 355 Suriname, Rombouts, H.E. 63 Suriname, Kramer, K.U., et al. 2745 Guyana, Anderson, C.W. 210 Guyana, Gillespie, L.J., et al. 1206 Suriname, Lanjouw, J. & J.C. Lindeman 2103 Guyana, Jansen-Jacobs, M.J., et al. 1567 Suriname, Berthoud-Coulon, M. 576 Bordenave, B. 816 Suriname, Elburg, J.P., LBB 13533 Mori, S.A., et al. 15223 Granville, J.J. de, et al. 9599 Boom, B.M., et al. 2507 Guyana, Singh 1 Petrov, I. 203 Guyana, Jenman, G.S. 7583 Guyana, Clarke, D., et al. 3013 Suriname, Lanjouw, J. & J.C. Lindeman 1873 Granville, J.J. de, et al. 9671 Guyana, Hahn, W.J., et al. 3895 Poncy, O., et al. 1874 Guyana, Little, E.L. 16949 Cremers, G., et al. 9332 Guyana, Henkel, T.W., et al. 3610 Benoist, R. 780 Richard, L.C.M. s.n. Mori, S.A., et al. 8794 Suriname, Mennega, A.M.W. 71 Guyana, Clarke, D., et al. 3476 Suriname, Splitgerber, F.L. 109 Prévost, M.F., et al. 2824 Hoff, M. 5087 Guyana, Clarke, D., et al. 3534 Guyana, Jansen-Jacobs, M.J., et al. 149 Hoff, M. 5090 Sabatier, D., et al. 2259 Suriname, Maguire, B. 23642 Guyana, Jenman, G.S. 7268 Suriname, Florschütz, P.A., et al. 238 Guyana, Gillespie, L.J., et al. 2249 Leblond, J.B. s.n. Suriname, Hostmann, F.W. & A. Kappler 1153 Granville, J.J. de, et al. 12581 Girod et al. 3237 Guyana, Forest Dept. British Guiana 4670 Granville, J.J. de, et al. 15272 Guyana, Talbot, H.F. s.n. Suriname, BW 6000 Suriname, Gonggrijp, J.W. 6594 Aublet, J.B.C.F. s.n. Suriname, Lanjouw, J. 701 Guyana, Henkel, T.W., et al. 3205 Granville, J.J. de, et al. 10958 Oldeman, R.A.A. T-269 Martin, J. 110 Grenand, P. 1306 Sastre, C., et al. 5761 Guyana, Clarke, D., et al. 7056 Guyana, Thurn, E.F. im s.n. Aublet, J.B.C.F. s.n. Desvaux, A.N. s.n. Prévost, M.F., et al. 1975 Guyana, Henkel, T.W., et al. 3177 Suriname, Lanjouw, J. & J.C. Lindeman 320 Service Forestier 4303 Suriname, Lim Sang 16250 Guyana, Henkel, T.W., et al. 2130 Guyana, Acevedo-Rodríguez, P., et al. 5812 Lescure, J.P. 272 Guyana, Jenman, G.S. 5333 Guyana, Forest Dept. British Guiana 6388 Suriname, Sterringa, J.T. LBB 12480 Suriname, Wullschlägel, H.R. 878 Guyana, Henkel, T.W., et al. 2110 Guyana, Henkel, T.W., et al. 4573 Prévost, M.F., et al. 2307 Guyana, Meyer, G. 113 Geay, F. 1905 Sagot, P.A. 944 Oldeman, R.A.A. 1225 Oldeman, R.A.A. 1371 Guyana, Hoffman, B., et al. 722 Hoff, M. 5982 Suriname, Lanjouw, J. & J.C. Lindeman 2853 Rohr, J.P.B. von 149 Guyana, Clarke, D., et al. 6191 Guyana, Forest Dept. British Guiana 4799 Suriname, Hostmann, F.W. & A. Kappler 812a Sastre, C., et al. 6364 Rohr, J.P.B. von 435 Oldeman, R.A.A. 1406 Guyana, Forest Dept. British Guiana 5457 Guyana, Forest Dept. British Guiana 3513 Guyana, Görts-van Rijn, A.R.A., et al. 485 Guyana, Hollowell, T.H., et al. 206 Mélinon, E.M. 381 Suriname, Versteeg, G.M. 647 Granville, J.J. de, et al. 6854 Guyana, Jenman, G.S. 6592 Guyana, Maas, P.J.M., et al. 4095 Guyana, Thurn, E.F. im s.n. Suriname, Vreden, C., LBB 16204 Guyana, Hoffman, B., et al. 1041 Guyana, Görts-van Rijn, A.R.A., et al. 9 Sabatier, D., et al. 2862 Suriname, Brownsbay 6594 Bordenave, B. 375 Billiet, F., et al. 1012 Loubry, D. 1215 Suriname, Rombouts, H.E. 833 Guyana, McDowell, T., et al. 2071 Guyana, Forest Dept. British Guiana 5690 Guyana, Cruz, J.S. de la 1141 Guyana, Clarke, D., et al. 1658 Service Forestier 7832 Granville, J.J. de, et al. 5768 Lemée, A.M.V. 1901 Guyana, Talbot, H.F. s.n. Martin, J. s.n. Richard, L.C.M. s.n. Skog, L.E., et al. 7456 Guyana, Hollowell, T.H., et al. 306 Suriname, Wullschlägel, H.R. s.n. Sagot, P.A. 240 Suriname, Splitgerber, F.L. 334 Oldeman, R.A.A. T-877 Mori, S.A., et al. 18653 Menten s.n. Suriname, Boerboom, J.H.A. 9140 Vahl, M. s.n. Suriname, Kock, C. s.n. Oldeman, R.A.A. 1403 Suriname, Stahel, G. 155 Suriname, BBS 91 Billiet, F., et al. 7492 Guyana, Forest Dept. British Guiana 4018 Loubry, D. 1934 Guyana, Henkel, T.W., et al. 3703 Guyana, Hoffman, B., et al. 3716 Sabatier, D., et al. 24 Barrier, S. 4901 Fleury, M. 446 Garnier, F.A. 109 Leprieur, F.R.M. s.n. Guyana, Pipoly, J.J., et al. 11247 Guyana, Acevedo-Rodríguez, P., et al. 3279 Suriname, Acevedo-Rodríguez, P., et al. 5812 Guyana, Watson, A.G. 31 Lemoult, E. 9 Prévost, M.F., et al. 4325 Lemoine, S.V. 7913 Suriname, Acevedo-Rodríguez, P., et al. 5954 Poiteau, P.A. s.n. Leblond, J.B. 83 Guyana, Pennington, T.D., et al. 410 Guyana, Jenman, G.S. 2168 Suriname, Heyde, N.M., et al. 89 Suriname, Lanjouw, J. & J.C. Lindeman 1912 Guyana, Anderson, C.W. 26 Granville, J.J. de, et al. 9614 Suriname, Focke, H.C. s.n. Béna, P. BAFOG 1313 Petrov, I. 208 Suriname, Outer, R.W. den 931 Guyana, Cruz, J.S. de la 3585 Guyana, Cruz, J.S. de la 4578 Oldeman, R.A.A. B-1926 Guyana, Acevedo-Rodríguez, P., et al. 3319 Martin, J. 450 Sastre, C., et al. 4117 Riera, B.J.J. 496 Guyana, Clarke, D., et al. 357 Suriname, Boon, H.A. 1257 Suriname, Florschütz, P.A., et al. 174 Suriname, Hostmann, F.W. & A. Kappler s.n. Suriname, Florschütz, P.A., et al. 161 Suriname, Hostmann, F.W. 1305 Sabatier, D., et al. 1134 Mori, S.A., et al. 22129 Granville, J.J. de, et al. 11889 Moretti, C. 824 Guyana, Hahn, W.J., et al. 4718 Guyana, Forest Dept. British Guiana 5770 Guyana, Bancroft, C.K., et al. s.n. Guyana, Hahn, W.J., et al. 4697 Guyana, Clarke, D., et al. 7737 Lescure, J.P. 122 Prévost, M.F., et al. 3648 Perrottet, G.S. s.n. Prévost, M.F., et al. 4895 Nolland, L.A. s.n. Suriname, BW 6261 Suriname, Hostmann, F.W. & A. Kappler 1032 Guyana, Smith, A.C. 2118 Guyana, Mutchnick, P., et al. 1359 Suriname, Evans, R., et al. 1910 Guyana, Jenman, G.S. 763 Granville, J.J. de, et al. 9296 Guyana, Rudge, E. s.n. Guyana, Gillespie, L.J., et al. 2253 Guyana, Forest Dept. British Guiana 3572 Suriname, Lindeman, J.C. & A.R.A. Görts et al. 350 Martin, J. s.n. Cremers, G., et al. 7192 Guyana, McDowell, T., et al. 3399 Guyana, Forest Dept. British Guiana 5810 Guyana, Maas, P.J.M., et al. 3619 Guyana, Polak, M., et al. 576 Granville, J.J. de, et al. B-4691 Oldeman, R.A.A. 2637 Granville, J.J. de, et al. T-1036 Mélinon, E.M. s.n. Suriname, Menge, J.G. s.n. Martin, J. 169 Guyana, Jansen-Jacobs, M.J., et al. 4951 Suriname, Schulz, J.P. 7221 Suriname, BW 3962 Guyana, Archer, W.A. 2373 Mélinon, E.M. 296 Perrottet, G.S. s.n. Service Forestier 7883 Guyana, Andel, T. van, et al. 1734 Leblond, J.B. 116 Oldeman, R.A.A. 3190 Feuillet, C. 1154 Patris, J.B. s.n. Suriname, Acevedo-Rodríguez, P., et al. 6006 Jacquemin, H. 2844 Suriname, Heyligers, P.C. 441 Guyana, Henkel, T.W., et al. 3730 Guyana, Hollowell, T.H., et al. 264 Guyana, Sandwith, N.Y. 1219 Cremers, G., et al. 7198 Guyana, King, K.F.S. 6399 Rohr, J.P.B. von 158 Guyana, Cruz, J.S. de la 1114 Sastre, C., et al. 8198 Suriname, Kappler, A. 1962 Guyana, Andel, T. van, et al. 1160 Suriname, Hulk, J.F. 382 Guyana, Cruz, J.S. de la 4129 Guyana, McDowell, T., et al. 1833 Guyana, Henkel, T.W., et al. 2709 Guyana, Stoffers, A.L., et al. 9 Oldeman, R.A.A. 972 Suriname, Lanjouw, J. & J.C. Lindeman 2076 Suriname, Splitgerber, F.L. s.n. Guyana, Hoffman, B., et al. 1031 Guyana, Jenman, G.S. 7800 Prévost, M.F., et al. 4550 Guyana, Cruz, J.S. de la 4156 Guyana, Mori, S.A., et al. 24347 Guyana, Clarke, D., et al. 3303 Guyana, Taylor 85 Suriname, Lanjouw, J. 815 Guyana, Maguire, B., et al. 23459 Prévost, M.F., et al. 1221 Sabatier, D., et al. 997 Lemoine, S.V. 7774 Suriname, BW 4085 Oldeman, R.A.A. 1791 Suriname, Mennega, A.M.W. 203 Suriname, Wessels Boer, J.G. 444 Guyana, Forest Dept. British Guiana 4221 Richard, L.C.M. s.n. Suriname, Heyde, N.M., et al. 647 Suriname, Lindeman, J.C. & A.L. Stoffers et al. 430 Suriname, Hugh-Jones, D.H. 47 Suriname, Daniëls, A.G.H. & F.P. Jonker 1326 Suriname, Heyde, N.M., et al. 273 Cremers, G., et al. 13739 Guyana, Henkel, T.W., et al. 2612 Guyana, Parker, C.S. s.n. Prévost, M.F., et al. 1359 Guyana, Cruz, J.S. de la 3074 Suriname, Hulk, J.F. 353 Guyana, Sandwith, N.Y. 475 Guyana, Clarke, D., et al. 1771 Guyana, McDowell, T., et al. 2135 Granville, J.J. de, et al. 77 Guyana, Forest Dept. British Guiana 6723 Guyana, Harrison, S.G. 1696 Guyana, Maas, P.J.M., et al. 3596 Sabatier, D., et al. 2300 Mélinon, E.M. 134 Suriname, BW 3343 Guyana, Forest Dept. British Guiana 4969 Suriname, Maas, P.J.M., et al. 2343 Guyana, Jansen-Jacobs, M.J., et al. 2473 Guyana, Hoffman, B., et al. 354 Prévost, M.F., et al. 1313 Guyana, Clarke, D., et al. 8875 Grenand, P. 3065 Le Goff, A. 63 Sauvain, M. 3 Guyana, Forest Dept. British Guiana 3580 Guyana, Jansen-Jacobs, M.J., et al. 1532 Guyana, Henkel, T.W., et al. 2047

Indumentum

Indumentum almost always of unicellular, slender, thick-walled, :33
33. 003

Inflorescences

Inflorescences , capitate to expanded, of simple or paniculate spikes or less often racemes;34
34. 004

Leaves

Leaves opposite (or whorled) or spiral (often ‘alternate’), petiolate, simple, entire, without stipules, often with a pair of petiolar glands.35
35. 002

Notes

A full synonymy for all taxa is given in .

Taxonomic changes

  • Combretum aubletii DC.

Taxonomy

Between 12 and 23 genera have been recognised in the past, the present author recognises 14 genera (Stace 2007a).
The genus Strephonema (West Africa) constitutes subfamily Strephonematoideae, the other 13 genera forming the Combretoideae. The latter is represented by 2 tribes, 1 of which has 2 subtribes; all 3 of these occur in the Guianas:
  • Laguncularieae: Laguncularia
  • Combreteae with 2 subtribes:
    • Combretinae: Combretum (including Thiloa)
    • Terminaliinae: Terminalia, Buchenavia and Conocarpus

Wood

Five Guianan genera are represented, 2 of which by 1 species only.
Due to the variation within the large genera Combretum and Terminalia, it is impossible to separate these genera. However, some characters can be used for a preliminary identification purposes. Noteworthy here is that the radial vessels reported by van Vliet (1979) could not be found in our material.
The parenchyma pattern is variable, even within species. According to van Vliet, the diagnostic value is restricted to those species “where distinctly different types (e.g. aliform and banded) are compared”.
A. -empty team- – In: InsideWood, B. Détienne, P. & J. Jacquet. 1983 – In: Atlas d’identification des bois de l’Amazonie et des régions voisines. – France: Centre Technique Forestier Tropical, C. Heimsch, C. 1942: Comparative anatomy of the secondary xylem in the Gruinales and Terebinthales of Wettstein with reference to taxonomic grouping. – Lilloa 8, D. IAWA Committee. 1989: The IAWA list of microscopic features for hardwood identification. – IAWA Bull. n.s. 10, E. Lens, F., S. Jansen, P. Caris, L. Serlet & E. Smets. 2005: Comparative wood anatomy of the primuloid clade (Ericales s.l.). – Syst. Bot. 30, F. Record, S.J. & R.W. Hess. 1936: Identification of woods with conspicuous rays. – Trop. Woods 48, G. Record, S.J. & R.W. Hess. 1943 – In: Timbers of the New World. – New Haven: Yale University Press, H. Vliet, G.L.C.M. van 1976: Radial vessels in rays. – IAWA Bull. 3, I. Vliet, G.L.C.M. van 1978: The vestured pits of Combretaceae and allied families. – Acta Bot. Neerl. 27, J. Vliet, G.L.C.M. van 1979: Wood anatomy of the Combretaceae. – Blumea 25