The Genera of Leguminosae-Caesalpinioideae and Swartzieae


L. Watson and M. J. Dallwitz

Character List

#1. ~ <Sensu lato genus to which sometimes referred (see Notes):>/

#2. <Synonyms:>/

An attempt has been made via this character to distinguish synonyms denoting (more or less) the same taxonomic concept as the name used here. It is also intended to encompass spelling variants of the ‘accepted’ name. Family names constituting nomenclaturally valid alternatives to those used are indicated (‘nom. altern.’). Combine characters 1 and 3 to list all the synonyms entered for a family.

‘Synonym’: a name rejected because of different taxonomic interpretation, or under the rules of nomenclature.

#3. Type species:/

Habit and leaf form

#4. <Habit:>/

1. trees/

2. shrubs/

3. herbs/

4. climbers or scramblers/

#5. <Presence of tendrils:>/

1. with tendrils/

2. without tendrils <implicit>/

#6. <Presence of specialised ‘short shoots’:>/

1. leaves and inflorescences crowded on ‘short shoots’ <or new shoots>/

2. without specialized ‘short shoots’/

#7. <Whether plants armed:>/

1. <plants> armed <with prickles and/or spines:>/

2. <plants> unarmed/

#8. <Whether exhibiting much reduced leaves:>/

1. exhibiting some leaves reduced to phyllodes, and/or with leaflets tiny and early-caducous/

2. without leaves reduced to phyllodes, the leaflets normally well developed and persistent <implicit>/

#9. Phyllotaxy/

1. distichous/

2. spiral/

#10. The leaves <overall form>/

1. compound/

2. ostensibly simple <or unifoliolate>/

3. <not compound but> two-lobed <or deeply emarginate>/

#11. The leaves <when simple or bilobed, venation>/

1. palmately veined <the midrib not predominant>/

2. with a predominant midrib/

#12. The leaves <mature compound, form>/

1. <mostly> pinnate <with three or more leaflets>/

2. <mostly> bipinnate/

3. bifoliolate <i.e., a compound leaf>/

#13. The leaves <bipinnate, insertion of pinnae>/

1. with <mostly> opposite or sub-opposite pinnae/

2. with alternate pinnae/

#14. The leaves <bipinnate, insertion of leaflets (pinnules)>/

1. with opposite or sub-opposite leaflets/

2. with alternate leaflets/

#15. The leaves <pinnation type>/

1. <commonly> imparipinnate <with well developed terminal leaflets: include trifoliate and digitate>/

2. <all> paripinnate <include unijugate>/

#16. The leaves <whether rachides are adaxially grooved (Polhill and Vidal 1981)>/

1. with adaxially grooved rachides <often with bridges or glands between the leaflet insertions>/

2. with rachides adaxially ridged <not grooved>/

#17. The leaflets <whether many or few per leaf>/

1. many per leaf <many leaves with eleven or more>/

2. few per leaf <most leaves with two to ten>/

#18. The leaflets <insertion in pinnate leaves>/

1. <mostly> opposite or sub-opposite/

2. <mostly> alternate/

#19. The leaflets <whether petiolulate>/

1. petiolulate/

2. sessile to sub-sessile/

#20. The leaflets <whether petiolules twisted (applicable only to pinnate leaves)>/

1. with markedly twisted petiolules/

2. without noticeably twisted petiolules/

#21. The <lateral> leaflets <symmetry>/

1. markedly asymmetrical <the lower half extending further down the midrib, and the laterals sometimes more or less palmate in its base; the upper half sometimes narrower, sometimes much reduced so that the displaced ‘midrib’ is almost marginal>/

2. symmetrical or nearly so <from their bases>/

#22. The leaflets <of bifoliolate, pinnate or bipinnate leaves, venation>/

1. palmately nerved <the midrib not predominant>/

2. with a predominant ‘midrib’ <this sometimes displaced to one side>/

#23. The leaflets <presence of a continuous marginal nerve>/

1. with a strong, continuous marginal nerve/

2. without a continuous marginal nerve/

This character has been awarded a low reliability for identification, because it is not reliably determinable from illustrations, and the source of the data recorded here is obscure. The present descriptions need checking against plant material.

#24. Stipules <presence, persistence (little information available on straightforward presence/absence)>/

1. present, persistent and <more or less> conspicuous in mature leaves/

2. absent or early caducous or very inconspicuous in mature leaves/

#25. Stipules <presence, form (better data required, for more precise character states)>/

1. leafy/

2. spinescent/

3. membranous <? - neither leafy nor spinescent>/

#26. Stipules <insertion>/

1. <more or less> connate/

2. not connate <separate, or joined only at the base>/

#27. Stipels <presence>/

1. present/

2. absent/


#28. The flowers <aggregated or solitary>/

1. aggregated into inflorescences <implicit>/

2. solitary/

#29. The inflorescences <position>/

1. axillary/

2. terminal/

#30. The inflorescences <simple or branched>/

1. unbranched/

2. branched/

#31. The inflorescences <branched, of racemose or cymose units>/

1. of racemose units/

2. of cymose units/

#32. The inflorescences <form>/

1. simple racemes/

2. simple corymbs/

3. fascicles <branched and congested in close clusters>/

4. panicles <indeterminate, open branching>/

#33. The flowers <insertion>/

1. distichous <in the inflorescence>/

2. not distichous <spiralled, or clustered>/

#34. Bracts <persistence (little information available on straightforward presence/absence)>/

1. absent at anthesis <i.e., early caducous or absent>/

2. persistent beyond anthesis/

#35. Bracteoles <presence>/

1. present/

2. absent/

#36. Bracteoles <form>/

1. small, not enclosing the flower buds/

2. relatively large and enclosing the flower buds/

#37. Bracteoles <when present, persistence>/

1. <present and> persistent beyond anthesis/

2. absent at anthesis <i.e., early caducous or absent altogether>/

#38. Bracteoles <aestivation>/

1. valvate/

2. <at least the lobes> not valvate <imbricate or distant>/

#39. Bracteoles <whether connate>/

1. connate/

2. free/


#40. The flowers <size/showiness>/

1. small/

2. large/

3. showy/

#41. The flowers <sexuality>/

1. <normally all> hermaphrodite/

2. <normally at least some> unisexual <should at least specify monoecious or dioecious ....>/

#42. The flowers <perianth and androecium: regular or zygomorphic>/

1. actinomorphic/

2. slightly irregular/

3. very irregular <including zygomorphic>/

#43. The flowers <mature, whether papilionaceous>/

1. ‘papilionaceous’ <exhibiting the full syndrome: see Notes>/

2. not <fully> ‘papilionaceous’/

‘Papilionaceous’: exhibiting a C5, imbricate-descending corolla as in typical Leguminosae-Papilionoideae, the butterfly shape deriving from an outer, adaxial (posterior) ‘standard’ petal (the vexillum), two lateral ‘wings’, and a ‘keel’ constituted by the anterior (lowermost, abaxial) pair; the latter enclosing the declinate androecium and gynoecium.

‘Pseudo-papilionaceous’: denotes a floral structure exhibiting some but not all of the features of, or superficially resembling, that typical of Leguminosae-Papilionoideae (q.v.).

#44. The flowers <symmetry>/

1. pentamerous <throughout K, C and A: K5, C5, A5, 10 or 15>/

2. not pentamerous throughout/

#45. The flowers departing from pentamery <whorls involved>/

1. in the calyx/

2. in the corolla/

3. in the androecium/

#46. The flowers <(calyx and/or corolla) colour>/

1. white or green/

2. <at least partially> coloured/

#47. Floral tube length relative to total hypanthium + calyx length <see Notes>/

Total length of ‘floral tube’ divided by tube + maximum calyx length, regardless of morphological interpretation.

#48. Hypanthium <presence: see Notes>/

1. present <the androecium perigynous>/

2. absent, the androecium hypogynous/

A saucer-, cup-shaped or tubular structure within which the the gynoecium is inserted, and on which the perianth and androecium are borne distally. Reasonably interpreted as representing evolutionary coalescence of the bases of perianth and androecium.

#49. Hypanthium <length relative to total flower length>/

1. very short/

2. short/

3. long/

4. very long/

#50. Hypanthium <shape>/

1. saucer-shaped/

2. cupular/

3. tubular/

4. campanulate/

#51. The perianth <presence/absence, constitution>/

1. comprising distinct calyx and corolla/

2. exclusively sepaline <more or less convincingly interpretable as sepals; ‘apetalous’>/

3. exclusively petaline <more or less convincingly interpretable as petals; ‘asepalous’>/

4. of ‘tepals’ <not readily resolvable into calyx and/or corolla>/

5. entirely vestigial/

6. absent <flower both asepalous and apetalous, achlamydeous>/

#52. Calyx <presence>/

1. present/

2. absent/

#53. Calyx <number of sepals or lobes>/

#54. Calyx <enclosure of flower buds>/

1. covering the rest of the flower in bud/

2. not covering the rest of the flower in bud/

#55. Calyx <whether Swartzieae type>/

1. Swartzieae type (closed before flowering, splitting more or less irregularly into valvate lobes or teeth)/

2. not Swartzieae type <implicit>/

#56. Calyx <coherence of members>/

1. gamosepalous <distal to the level of insertion of the corolla>/

2. polysepalous <includes ‘segments separate to the hypanthium’>/

#57. Calyx <symmetry>/

1. more or less regular <the segments more or less equal>/

2. markedly irregular <the segments very unequal; includes bilabiate>/

#58. Calyx members <aestivation>/

1. imbricate <in the bud>/

2. not imbricate <valvate or sub-valvate, or the segments separate in bud>/

#59. Corolla <presence>/

1. present <implicit: including any minute petals>/

2. absent/

#60. Corolla <symmetry>/

1. regular <actinomorphic>/

2. slightly irregular/

3. very irregular/

#61. Corolla <number of petals or lobes, including any tiny members>/

#62. Corolla <whether incorporating tiny or vestigial members>/

1. including greatly reduced <tiny or more or less vestigial> members/

2. without greatly reduced <tiny or vestigial> members/

#63. Well developed petals <number, location>/

#64. Vestigial petals <number, location>/

#65. Corolla <coherence of members>/

1. gamopetalous/

2. polypetalous/

3. monopetalous/

#66. Petals <clawed, presence>/

1. clawed <i.e., at least one>/

2. sessile <not clawed>/

#67. Petals <aestivation in bud>/

1. imbricate/

2. valvate/

#68. Petals <direction of aestivation>/

1. imbricate-ascending <the posterior petal (vexillum) inside>/

2. imbricate-descending with the posterior petal (vexillum) outside <including but not exclusively comprising ‘papilionaceous’ forms>/

#69. Petals <colour (/

1. white <or greenish>/

2. yellow <to orange>/

3. <pink to> red <to purple>/

#70. Disk <presence>/

1. present and conspicuous/

2. absent <or very inconspicuous>/

#71. The androecium comprising <number of members, including any staminodes>/


#72. The androecium <peculiar staminal morphology>/

1. characterized by the long, sigmoidally incurved filaments of the three abaxial stamens being many times longer than their small anthers, which are dorsifixed, sub-versatile and introrsely dehiscent via short slits, contrasting with the short adaxial stamens with their straight filaments and anthers dehiscing via basal pores (Cassia)/

2. without the three long-filamented, short-anthered abaxial stamens characteristic of Cassia/

#73. The androecium <whether declinate>/

1. declinate/

2. not declinate/

#74. The androecium <coherence of parts>/

1. with united members <i.e., at least some united at least at the base; include any staminodes>/

2. members all free of one another/

#75. The androecium members <relative lengths of stamens and any staminodes>/

1. all more or less equal in length/

2. markedly unequal/

#76. The androecium <presence of staminodia (information lacking on numbers of staminodia)>/

1. including staminodia/

2. comprising only fertile stamens <without staminodia>/

#77. The staminodia <number>/

#78. The staminodia <description>/

#79. Fertile stamens <number; ‘many’ arbitrarily scored as ‘26 or more’>/

#80. Anthers <position of attachment to filament>/

1. attached at the base of the connective/

2. attached well above the base of the connective <implies anthers versatile>/

#81. Anthers dehiscing <orientation>/

1. introrsely/

2. laterally/

#82. Anthers dehiscing <by pores or slits>/

1. <at least initially> via pores or short slits/

2. longitudinally/

#83. Ovary <insertion>/

1. sessile or subsessile/

2. stipitate/

#84. Ovary <adnation>/

1. free/

2. eccentric, with the stipe adnate <stipe more or less adnate to the receptacle, calyx or hypanthium>/

#85. Stigma <whether dilated (information for precise character states unavailable)>/

1. dilated <expanded: capitate, peltate or sub-peltate>/

2. not dilated <truncate, oblique, punctiform: implicit>/

#86. Ovules <number (insufficient information available to record as a numeric character)>/

1. solitary/

2. few <2–5>/

3. numerous <more than 5>/

Fruit, seed and seedling

#87. Fruit <type>/

1. a follicle/

2. a two-valved pod/

3. indehiscent/

#88. Fruit <whether drupaceous>/

1. drupaceous/

2. not drupaceous <implicit>/

#89. Fruit <shape: whether straight or curved>/

1. <more or less> straight/

2. <distinctly> curved/

#90. Fruit <whether internally septate>/

1. internally septate <the seeds separated>/

2. not internally septate/

#91. Fruit <whether the valves twist on dehiscence>/

1. valves twisting and enrolling during dehiscence/

2. without markedly twisting or enrolling valves/

#92. Fruit <wings, location>/

1. winged longitudinally/

2. winged distally to the single seed/

3. winged proximally to the single seed/

4. broadly papery-winged all round/

5. not noticeably winged <implicit>/

#93. Fruit <number of wings>/


#94. Fruit <texture>/

1. becoming woody/

2. not becoming woody <leathery, fleshy, membranous etc.>/

#95. The mature <fruit> valves <venation, ignoring sutures>/

1. with conspicuous, prominent, raised venation/

2. without prominent venation/

#96. The conspicuous <fruit valve> venation <alignment at maturity>/

1. predominantly longitudinal/

2. not predominantly longitudinal <oblique, transverse or reticulate>/

#97. Seeds <whether endospermic>/

1. endospermic/

2. non-endospermic/

#98. Seeds <whether arillate>/

1. arillate/

2. not arillate/

#99. Seeds <radicle shape>/

1. with a straight or slightly oblique radicle/

2. with an inflexed radicle/

#100. Seeds <amyloid in cotyledons and/or any endosperm (Kooiman 1960)>/

1. amyloid-positive/

2. amyloid-negative/

#101. Seeds <(amyloid-negative) whether with galactomannan or starch (Kooiman 1960)>/

1. with galactomannan/

2. with starch/

#102. Cotyledons <form, texture>/

1. <‘foliaceous’ or> flat/

2. <not ‘foliaceous’,> not flat <fleshy, folded, wrinkled etc.>/

#103. Cotyledons of <type (Smith 1981)>/

1. Type 1 <leaflike, folded in seed, expanding on germination, with abundant palisade>/

2. Type 2 <leaflike, flat in seed, expanding on germination, with abundant palisade>/

3. Type 3 <fleshy storage organs, with little or no expansion on germination; some imprecisely demarcated palisade>/

4. Type 4 <fleshy storage organs, with no expansion on germination; no palisade>/

#104. Cotyledons <(fleshy), vascular arrangement (Smith 1981)>/

1. with a vascular system in one plane/

2. with a vascular system ramified throughout/

#105. Cotyledons <germination type>/

1. epigeal/

2. hypogeal/

Transverse section of lamina

#106. Leaves <presence of phloem transfer cells (Watson and Gunning 1981)>/

1. with <more or less> conspicuous phloem transfer cells in the minor veins/

2. without conspicuous phloem transfer cells in the minor veins/

#107. Druses <presence in the mesophyll>/

1. common in the mesophyll/

2. absent from <or very rare in> the mesophyll/

#108. Mesophyll secretory cavities <presence>/

1. <which are usually visible as> (gland-dots) common/

2. absent <or very rare, i.e., leaves not gland-dotted>/

#109. Mesophyll secretory cavities <type>/

1. epithelium-lined/

2. without a lining of epithelium/

#110. Adaxial hypodermis <presence>/

1. present/

2. absent/

#111. Leaf girders <sclerenchymatous or fibrous, occurrence>/

1. common (the veins transcurrent)/

2. absent <or very rare, the veins ‘embedded’: ignore midribs of leaves and leaflets>/

#112. Laminae <mesophyll organisation>/

1. isobilateral, with adaxial and abaxial palisades/

2. dorsiventral/

#113. Mesophyll <presence of unaligned fibres or sclereids>/

1. exhibiting fibres or sclereids which are unaligned with the vascular bundles/

2. without unaligned fibres or sclereids/

#114. Minor veins <fibre accompaniment>/

1. mainly with abundant accompanying fibres/

2. lacking accompanying fibrous tissue/

Leaf lamina epidermes

#115. Epidermal crystals <presence>/

1. present <adaxially and/or abaxially>/

2. not seen either adaxially or abaxially/

#116. Epidermal crystals <form>/

1. prisms/

2. druses/

#117. Simple unbranched hairs <presence>/

1. common <adaxially and/or abaxially on the laminae>/

2. not seen/

#118. Simple unbranched hairs <surface texture>/

1. scabrid/

2. smooth/

#119. <Presence on the laminae of compound or branched eglandular hairs>/

1. compound or branched eglandular hairs present/

2. no compound or branched eglandular hairs seen/

#120. <Presence on the laminae of peltate or> capitate glands/

1. present/

2. not seen/

#121. Hooked hairs <presence on the laminae>/

1. present <adaxially and/or abaxially>/

2. not seen/

#122. Cassieae-type leaf pseudo-glands <presence on the laminae>/

1. present <adaxially and/or abaxially>/

2. not seen/

#123. Expanded and embedded hair-feet <presence on the laminae>/

1. present <adaxially and/or abaxially>/

2. absent/

#124. Expanded and embedded hair-feet <wall structure>/

1. all medium to thick-walled/

2. (at least some of them) thin-walled/

#125. Expanded and embedded hair feet <complexity>/

1. (at least some of them) complex with vertical walls/

2. all simple, without vertical walls/

#126. Basally bent hairs <presence on the laminae>/

1. present <abaxially and/or adaxially>/

2. absent/

#127. Adaxial interveinal epidermal cell walls <shape>/

1. <more or less> straight <or gently undulating> in optical section/

2. markedly sinuous <or tessellated, at least> in high-focus optical section/

#128. Adaxial interveinal epidermal cell walls <whether pitted>/

1. conspicuously pitted <in optical section>/

2. not conspicuously pitted <in optical section>/

#129. Adaxial interveinal epidermal cell walls <thickness in outer optical section>/

1. thick/

2. medium-thick/

3. of medium thickness/

4. medium-thin/

5. thin/

#130. Stomata <adaxial, occurrence>/

1. adaxially common and widespread/

2. adaxially <absent or> very rare <or very localised, e.g., only near main veins>/

#131. Abaxial stomata <form>/

1. <ignoring those over veins, and giants> predominantly paracytic/

2. not predominantly paracytic <actinocytic, cyclocytic, anomocytic or a mixture>/

#132. Abaxial epidermis <papillation>/

1. papillate interveinally/

2. not papillate <interveinally>/

#133. Abaxial epidermis <whether papillae over-arch or obscure the stomata>/

1. with papillae <more or less> over-arching <or obscuring> the stomata/

2. with the papillae not over-arching the stomata/

#134. Abaxial interveinal epidermal cell walls <shape>/

1. straight, or gently undulating/

2. markedly sinuous <or tessellated, at least> in high-focus optical section/

#135. Abaxial interveinal epidermal cell walls <whether pitted>/

1. conspicuously pitted in optical section/

2. not conspicuously pitted in optical section/

#136. Abaxial interveinal epidermal cell walls <affinity for safranin>/

1. scarcely staining with safranin/

2. staining normally with safranin/

#137. Abaxial interveinal epidermal cell walls <thickness in outer optical section>/

1. thick/

2. medium-thick/

3. of medium thickness/

4. medium-thin/

5. thin/

Wood anatomy

#138. Wood <presence of septate fibres>/

1. with septate fibres/

2. without septate fibres/

#139. Wood <organisation>/

1. <at least partially> storied/

2. not storied/

#142. Intervascular pits <size>/

1. very small <3 microns, or less>/

2. medium to large <more than 3 microns>/

Pollen ultrastructure

#143. Tectum <of pollen grains, general surface pattern>/

1. punctate/

2. reticulate/

3. striate/

4. papillate/

#144. Tectum <of pollen grains, punctation type>/

1. smooth punctate, or puncticulate/

2. rugulose punctate/

#145. Tectum <of pollen grains, reticulation type>/

1. finely to moderately regularly reticulate/

2. strongly irregularly coarse-reticulate/

3. rugulose reticulate/

4. verrucose reticulate/

#146. Tectum <of pollen grains, striation type>/

1. pole to pole striate/

2. reticulate striate/

3. interwoven striate/

#147. Tectum <of pollen grains, papillation type>/

1. simple papillate <i.e., not reticulate papillate>/

2. reticulate papillate/

#148. Length of colpi/

1. greater than one half pole to pole distance/

2. less than one half pole to pole distance/

#149. Foot layer of pollen wall <texture>/

1. smooth/

2. with obvious projections/


#150. Basic chromosome number, x = <Goldblatt 1981>/

#151. 2n = <diploid chromosome number: data from Adv. Leg. Syst. 1981>/

Species number and distribution.

#152. <Number of species:>/


#153. <World distribution:>/

1. Western Eurasia, U.S.S.R. <includes Iran, Iraq, Turkey>/

2. Mediterranean/

3. Eastern Asia <Japan, China to India>/

4. Africa <and Saudi Arabia>/

5. Madagascar/

6. Pacific <Malaysia, Indonesia, Australasia, Pacific islands>/

7. North America <U.S.A., Mexico>/

8. South and Central America/

9. West Indies/

This character is intended for use only in pursuing identifications. The very broadly defined regions are without phytogeographical import.

#154. <Geographical distribution:>/


#155. <Tribe:>/

1. Caesalpinieae/

2. Cassieae/

3. Cercideae/

4. Detarieae <Amherstieae of Cowan and Polhill (1981) indicated in the descriptions>/

5. Papilionoideae-Swartzieae/

6. Papilionoideae-Amburaneae (former Swartzieae)/

7. Papilionoideae-Exostyleae (former Swartzieae)/

#156. <Clade within Detarieae (Bruneau et al., 2008):>/

1. Amherstieae clade of Bruneau et al. (2008)/

2. Prioria clade of Bruneau et al. (2008)/

3. Detarieae sensu stricto clade of Bruneau et al. (2008)/

Economic uses, comments

#157. <Economic uses, etc.>/

#158. <Whether widely cultivated (preliminary data only; information on economic uses not yet compiled):>/

1. widely cultivated/

2. not widely cultivated /

#159. <Unusual features characteristic of the genus:>/

#160. <General comments:>/


#161. Abbreviated taxon name:/

#162. Illustrations:/

We advise against extracting comparative information from the descriptions. This is much more easily achieved using the DELTA data files or the interactive key, which allows access to the character list, illustrations, full and partial descriptions, diagnostic descriptions, differences and similarities between taxa, lists of taxa exhibiting or lacking specified attributes, distributions of character states within any set of taxa, geographical distribution, and classification. See also Guidelines for using data taken from Web publications.

Cite this publication as: ‘Watson, L., and Dallwitz, M.J. 1993 onwards. The genera of Leguminosae-Caesalpinioideae and Swartzieae: descriptions, illustrations, identification, and information retrieval. In English and French. Version: 22nd March 2017.’.