Hererolandia most closely resembles Lophocarpinia, but differs in having scattered curved, deflexed prickles on shoots (vs. scattered straight, conical spines, as well as modified, short, lateral, spinescent branchlets), pinnate leaves with (4–) 5–7 (–9) pairs of leaflets, arranged in fascicles (vs. alternate, pinnate leaves with 2–3 pairs of leaflets), and leaflets elliptic to oblong-elliptic (vs. leaflets obovate or elliptic-orbicular). The most distinctive feature of Hererolandia is the thinly woody, laterally compressed, almost circular to strongly sickle-shaped, usually 1-seeded fruit, covered in robust trichomes up to 6 mm long (vs. a segmented, falcate, lomentaceous fruit, with 4 coarsely serrate wings, breaking up into 1-seeded units).

Hererolandia pearsonii (L. Bolus) E. Gagnon & G. P. Lewis ≡ Caesalpinia pearsonii L. Bolus

A multi-stemmed shrub to 2 m, but usually less than 1 m tall, armed with curved, deflexed, 7 mm long prickles scattered along the branches; bark white or brown; stems terete and slightly sinuous, with a fine silvery indumentum on the young twigs, older stems glabrescent. Stipules not seen. Leaves pinnate, 7–17 mm long, subsessile, borne in fascicles on short woody brachyblasts that are usually subtended by a pair of tiny (sometimes obscure) prickles; leaflets opposite, (4–) 5–7 (–9) pairs per pinna, eglandular, covered in a fine silvery pubescence, 5–6.5 × 2.5–3 mm, elliptic to oblong-elliptic, apex obtuse, with an acuminate tip, main vein prominent, secondary venation not visible. Inflorescence a short raceme of bisexual flowers, about 5 cm long, usually borne on brachyblasts, covered in a fine silvery pubescence, with prickles along the inflorescence rachis; bracts about 2–3 × 1.5 mm, ovate, apex acute, caducous. Flowers zygomorphic; calyx with a short hypanthium, and 5 free sepals, c. 3–5 mm long, finely white pubescent, with the lower sepal cucullate and covering the other 4 sepals in bud, all sepals caducous, but hypanthium persistent as a ring around the stipe of the fruit; petals 5, yellow, free, c. 6–9 mm long, obovate; stamens 10, free, up to 10 mm long, eglandular, pubescent on the lower half; ovary pubescent, stigma a fringed and slightly indented chamber. Fruit a thinly woody, laterally compressed, almost circular to strongly sickle-shaped pod, c. 2–2.3 × 1–1.5 cm, dehiscing along the sutures, finely pubescent and covered in robust trichomes up to 6 mm long, usually 1-seeded. Seeds laterally compressed, about 6–8 mm long.

A monospecific genus endemic to Namibia, on the Great Escarpment.

Semi-desert and desert areas, on stony, sandy soils.

Semiarid Hereroland, a region of eastern Namibia, is the type locality of H. pearsonii. The Herero people who inhabit this region are nomadic cattle herders and it is they and their region that are honoured in the name proposed for this monospecific genus, endemic to this restricted area of Namibia.

Bolus (1920); Roux (2003); Curtis and Mannheimer (2005: 227).

Figure 4. 

Hererolandia pearsonii (L. Bolus) E. Gagnon & G. P. Lewis. A foliage and inflorescences B stem armature detail C leaflet lower surface D calyx lobes outer surface E lower cucullate calyx lobe side view F median petal inner surface G median petal side view H upper lateral petal inner surface I lower lateral petal inner surface J stamens and part of gynoecium, with calyx lobes removed K anthers dorsal and ventral views L gynoecium M stigma detail, N fruit. A, C–M from Müller 1006, B, N from Geiss et al. 5156. Drawn by Juliet Williamson.

Figure 5. 

Hererolandia pearsonii (L. Bolus) E. Gagnon & G. P. Lewis. A shrubby habit B inflorescence C branch showing prickles and leaves D fruits (A. A. Dreyer, Sesriem Canyon, Namibia, unvouchered). Haematoxylum brasiletto H. Karst. E mature fruit dehiscing along the mid-valve (C. E. Hughes, Mexico, unvouchered) F inflorescences and leaves (G. P. Lewis, Mexico, Lewis 2057 (K)) G distinctively fluted trunks (C. E. Hughes, Oaxaca, Mexico, Hughes 1947 (FHO)) Lophocarpinia aculeatifolia (Burkart) Burkart H shrub with flowers, armed with straight conical spines I fruits (R. H. Fortunato, Paraguay, Fortunato 8650 (BAB)).

Caesalpinia pearsonii L. Bolus, Annals of the Bolus Herbarium 3: 4. 1920.

NAMIBIA, Ababes, breccia banks of Tsondab River below farm, 29 Dec 1915, Pearson 9162 (holotype: BOL; isotypes: K!, GRA, NBG, PRE).

Lophocarpinia aculeatifolia (Burkart) Burkart ≡ Cenostigma aculeatifolium Burkart.

Shrub 0.5 (– 3) m tall, armed with scattered straight, conical, 2–5 mm long spines on shoots; leaves and inflorescences crowded on brachyblasts; shoots glabrous, reddish, the lateral ones sometimes, spinescent. Stipules acuminate, caducous. Leaves alternate, paripinnate, 5–10 mm long; leaflets in 2 (– 3) pairs, obovate or elliptic-orbicular, 4–7 × 2–2.4 mm, finely pubescent, eglandular, with a pair of small prickles at the insertions of the leaflets. Inflorescences short, corymbiform, pubescent racemes, each with 3–6 bisexual flowers; bracts small, caducous. Flowers zygomorphic, 1–1.5 cm long; calyx with a turbinate, fleshy hypanthium, and 5 oblong, pubescent, caducous sepals, lower sepal cucullate and covering the other 4 sepals in bud, embracing the androecium and gynoecium at anthesis; petals 5, yellow to yellow-orange, free, the median petal differentiated from the rest by a fleshy claw and wavy blade margins, pubescent; stamens 10, free, filaments pubescent; ovary glabrous; stigma apical, concave. Fruit a lomentum, with 1–5 segments, falcate, with 4 coarsely serrate wings. Seeds ellipsoid to reniform, smooth.

A monospecific genus restricted to Argentina and Paraguay (possibly also occurring in Mato Grosso do Sul, Brazil, pers. comm. H. C. de Lima).

Chaco woodland and seasonally dry tropical to subtropical forest.

From lopho- (Greek: combed or crested) and carpos (Greek: fruit), the fruit has 4 crested wings, the ending -inia signifies a close relationship with Caesalpinia.

Burkart (1957); Ulibarri (2008); Nores et al. (2012).

Figure 6. 

Lophocarpinia aculeatifolia (Burkart) Burkart. A habit B flowering and fruiting branch C detail of leaf attachment D, E median petal front and side views F upper lateral petal G lower lateral petal H stamen I gynoecium J stigma K fruit longitudinal section L fruit cross section. A, B from Burkart 20216 C, K, L after illustration by BurkartD–J from Burkart 20218. Drawn by Christi A. Sobel.

Haematoxylum campechianum L.

Multi-stemmed shrubs to 3 m, to medium-sized trees, 3–15 m in height, armed with scattered straight conical spines, 0.5–1.5 cm long on shoots, and the short, lateral shoots spinescent; mature trees with conspicuously fluted trunks, shrubs often with ribbed branches; young stems reddish brown to grey, glabrous to pubescent, eglandular (or with stalked glands in H. dinteri). Leaves alternate, pinnate or bipinnate (both can be present on the same individual in some species), glabrous to pubescent, eglandular, 1–10 cm long; pinnate leaves with 2–6 pairs of leaflets, 2.5–35 × 3–30 mm, glabrous to slightly pubescent, eglandular; bipinnate leaves with 1–3 pairs of pinnae plus a terminal pinna, each pinna with 2–5 (–6) pairs of leaflets, 5–11 × 2–4.5 mm; leaflets in opposite pairs, obcordate to obovate, apex emarginate to obtuse, base cuneate to attenuate (occasionally obtuse), short-petiolulate; primary vein centric, secondary veins ascending, and forming a sharp angle with the primary vein. Inflorescences terminal or axillary racemes or panicles of pedicellate flowers; rachis and pedicels unarmed, glabrous to pubescent, eglandular or glandular. Flowers bisexual, actinomorphic to zygomorphic; calyx comprising a hypanthium and 5 free sepals that are c. 6–7 mm long, glabrous to pubescent, eglandular or glandular, the lower sepal cucullate and slightly covering the other 4 in bud, sepals caducous, hypanthium persisting in fruit, forming a calyx ring; petals 5, yellow to pale yellow or white, free, imbricate, obovate to oblanceolate, 4–10 mm long; stamens 10, free, filaments pubescent, particularly on the lower half; ovary glabrous to pubescent. Fruit flattened, membranaceous to chartaceous, oblong to fusiform (occasionally falcate), apex rounded to obtuse, base acute, dehiscing along the middle of the valves, or near the margin of the fruit, but never along the sutures, 10–50 × 4–15 mm, 1–3-seeded. Seeds oblong to reniform, flattened, 6–12 × 3.8–5 mm.

Haematoxylum comprises five species: two in Central America (Salvador to Costa Rica), Mexico, South America (Colombia and Venezuela) and the Caribbean (perhaps introduced), two endemic to Mexico and one in Southern Africa (Namibia).

Deserts, seasonally dry tropical semi-deciduous scrub and thorn scrub, sandy river beds and dry rocky hillsides. One species (H. campechianum) is known to grow in frequently inundated marshy areas by rivers.

From haemato- (Greek: bloody) and xylon (Greek: wood), alluding to the blood-red heartwood of H. campechianum L. which produces a brilliant red dye.

There is a key to species by Durán and Sousa, in Novon 23(1): 31–36 (2014).

Standley and Steyermark (1946); Ross (1977: 122–114); Roux (2003); Curtis and Mannheimer (2005: 215); Durán and Ramírez (2008); Barreto Valdés (2013); Durán and Sousa (2014).

Figure 7. 

Haematoxylum campechianum L. A flowering branch B leaflet C flower bud D flower E median petal F lateral petal G stamen H gynoecium I infructescence J dehiscing fruit K seed L embryo. A, B, D–H from Lorence 2746 C from Balfour s.n. I–L from Johnston s.n. Drawn by Eleanor Catherine, originally published in Flore des Mascareignes 80. Légumineuses, page 6, plate 1.

Paubrasilia is closely related to Caesalpinia, but differs in habit, forming medium-sized to large trees, 5–15+ m tall, armed with small to large upturned prickles, these usually arising from woody protuberances (vs. shrubs or small to medium sized trees, usually 1–6 m tall, unarmed or armed with curved deflexed prickles, either occurring in pairs at the base of leaves, or scattered on shoots, or both, and sometimes present at the base of trunk). Paubrasilia also differs from Caesalpinia by having alternate pinnae with consistently alternate leaflets (vs. opposite pinnae with opposite to alternate leaflets), the median petal with a blood red central blotch (vs. the median petal lacking a red central blotch) and a spiny, woody, finely pubescent, sub-lunate, 1–2-seeded pod (vs. an unarmed, glabrous, oblong-elliptic, generally 3–7-seeded pod, with a marcescent style forming an acute apex).

Paubrasilia echinata (Lam.) E. Gagnon, H.C. Lima & G. P. Lewis ≡ Caesalpinia echinata Lam.

Medium sized to large trees, 5–15+ m tall, armed with small to large upturned prickles, these usually arising from woody protuberances, 1–20 mm long (the prickles often sparse or lacking on more mature specimens and larger, older branches); bark chestnut brown to almost black with greyish pustular lenticels, flaking in large woody plates; heartwood red, with the trunk exuding a red sap when injured. Stipules lanceloate, acute to acuminate, caducous. Leaves bipinnate, ending with a pair of pinnae; petiole and rachis finely tomentose; pinnae alternate, the terminal pair opposite to subopposite, with (2–) 3–20 pairs of pinnae per leaf; leaflets alternate, with (2–) 3–19 (–21) leaflets per pinna (generally the number of leaflets is inversely proportional to their size), 0.9–5 × 0.5–3.6 cm (although some specimens have leaflets up to 12 cm long), leaflet blades coriaceous, broadly oblong to subrhombic, apex rounded, obtuse or emarginate, base asymmetric, eglandular, glabrous, midvein excentric, secondary veins brochidodromous. Inflorescence a terminal, or occasionally axillary, finely tomentose raceme or panicle, with c. 15–40 flowers; bracts broadly ovate-triangular, apex acute to acuminate, less than 1 mm long, pubescent, caducous. Flowers bisexual, zygomorphic; calyx a tomentose hypanthium with 5 sepals, that are c. 5–9 mm long, the lowest sepal cucullate, covering the other 4 in bud, all sepals caducous but the hypanthium persisting as a free ring around the pedicel as the pod matures; petals 5, free, bright yellow, the median petal with a blood-red blotch on the inner face, c. 11–15 × 4–10 mm, all petals eglandular, broadly-obovate to slightly spathulate, the petal claws pubescent; stamens 10, free, 7–9 mm long, eglandular, densely pubescent on lower half; ovary pubescent with small spines intermixed, stigma a subterminal fringed-chamber. Fruit a spiny, finely pubescent, sub-lunate, woody, 5.5–7.3 × 1.9–2.6 cm, elastically dehiscent pod with twisting valves, 1–2-seeded. Seeds laterally compressed, ovate-obovate.

A monospecific genus endemic to Eastern Brazil, in the states of Pernambuco, Bahia, Espirito Santo and Rio de Janeiro. Widely cultivated in Brazil as an ornamental street or park tree, and sometimes in plantations.

Dry coastal cactus scrub often on rocky outcrops, inland in Mata Atlântica, and in tall restinga on well-drained sandy soil.

“Pau-brasil” is the national tree of Brazil, and has long been associated with the country. Its red sap was once used for dying cotton and cloth and its wood is much prized for the manufacture of high quality violin bows. Originally described as Caesalpinia echinata by Lamarck in 1785, it is appropriate that this phylogenetically isolated taxon should be placed in its own monospecific genus and a Latinization of its well-known and much used common name recognises the importance of the species to Brazil. For a detailed account of this iconic species refer to Pau-brasil by E. Bueno [et al.], São Paulo, Axis Mundi (2002).

Lewis (1998: 152–158); Bueno (2002); Cardoso et al. (2005).

Figure 8. 

Paubrasilia echinata (Lam.) E. Gagnon, H. C. Lima & G. P. Lewis. A inflorescences and foliage B leaflet undersurface C bark armature (front and side views) D flower E flower l.s. F median petal G upper lateral petal H lower lateral petal I stamen J gynoecium K stigma L fruit M single valve of dehisced fruit N seedling. A from Glaziou 6839 B, K from Angeli 201 C, M from Lewis et al. 1634 D from Lima et al. 2705 E–J from Ducke 20623 L from Mell s.n., N from Lewis et al. 1624. Drawn by Tim Galloway.

Figure 9. 

Paubrasilia echinata (Lam.) E. Gagnon, H. C. Lima & G. P. Lewis. A flowers (H.C. Lima, Brazil, Lima et al. 2705 (RB)) B fruits (G. P. Lewis, Brazil, unvouchered) C prickles on woody protuberances on a young trunk (E. Gagnon, Bahia, Brazil, Lima et al. 7909 (RB)) D habit (L. P. de Queiroz, Bahia, Brazil, unvouchered) E fluted trunk of a mature individual (E. Gagnon, Bahia, Brazil, Lima et al. 7894 (RB) F cross section of the trunk, showing dark red heartwood (E. Gagnon, Espirito Santo, Brazil, unvouchered), G inflorescences (L. P. de Queiroz, Bahia, Brazil, unvouchered).

Caesalpinia echinata Lam., Encycl. 1: 461. 1785. Guilandina echinata (Lam.) Spreng., Syst. Veg. 2: 327. 1825.

[BRAZIL] “In locis mari vicinis non apparet, sed tantum in mediterraneis silvis, unde magno labore ad littoralia vehitur” (Lectotype: [icon] “Ibirapitanga, sive Lignvm Rvbrvm” in Piso, De Indiae utriusque re naturali et medica: 164. 1658, designated here).

An epitype is to be selected in a subsequent paper focussing on the morphotypes of P. echinata (De Lima et al., in prep.).

Caesalpinia vesicaria Vell., Fl. Flumin.: 172. 1829, Fl. Flumin. Icon. 4. t. 89. 1831. (“vessicaria”), non L. 1753. .

Type. [BRAZIL], “Habitat silvis maritimis usque ad Molendinum Sacchariferum dictum Itacurussá” (Lectotype: [icon] “Cæsalpinia vessicaria” in Velloso, Fl. Flumin. Icon. 4: t. 89. 1831).

Caesalpinia obliqua Vogel in Linnaea 11: 407. 1837.

Type: BRAZIL, Sellow s.n. (holotype ? B †; isotype P02142646!).

Caesalpinia resembles Guilandina, but differs in habit, comprising armed shrubs and small trees (vs. armed lianas and scrambling/trailing shrubs). It also differs in having racemes of bisexual flowers (vs. racemes of unisexual flowers), sepals imbricate in bud, with a pronounced lower cucullate sepal (vs. sepals valvate in bud), zygomorphic corollas variable in colour (yellow, white, red, orange, pink or green), with petals extending well beyond the sepals (vs. zygomorphic to sub-actinomorphic, yellow corollas, with petals barely extending beyond the sepals), coriaceous, oblong-elliptic to linear, laterally compressed, glabrous pods (vs. oblong-elliptic inflated pods, usually armed with 5–10 mm long spinescent bristles), and obovoid, laterally compressed seeds (vs. obovoid globular seeds).

Caesalpinia brasiliensis L.

Shrubs or small trees, usually 1–6 m tall, armed with curved deflexed prickles (except C. nipensis which is unarmed), these either in pairs at the base of leaves, or scattered along the shoots (or both), or sometimes on woody protuberances at the base of trunks and stems; young shoots terete, glabrous and eglandular. Stipules not seen. Leaves alternate, bipinnate, c. 4–30 cm long, ending with a pair of pinnae, unarmed, or sometimes with a pair of prickles at the insertion of the pinnae on the leaf rachis, sometimes also at the insertions of the leaflets on the pinna rachis; pinnae opposite, in (1–) 2–6 pairs per leaf; leaflets alternate to opposite, in 3–13 pairs per pinna, short-petiolulate, blades suborbicular, obovate or elliptic, apex mucronate, rounded or emarginate, base cuneiform, rounded or oblique; main vein centric, secondary veins reticulate. Inflorescence a terminal or axillary raceme or panicle of pedicellate, bisexual flowers, c. 5–37 cm long, unarmed; bracts lanceolate or ovate, apex acute to acuminate, caducous. Flowers zygomorphic, c. 13–25 mm long; calyx comprising a hypanthium with 5 sepals, that are each c. 7–17 mm long, glabrous to occasionally finely puberulous, always eglandular, the lower sepal strongly cucullate and covering the other 4 sepals in bud, all sepals caducous, but hypanthium persistent as a free ring around the pedicel as the fruit matures; petals 5, variable in colour (yellow, white, red, orange, or green; certain horticultural varieties are also pink), the corolla also variable in shape (related to different pollination systems: bees, butterflies, birds and bats); stamens 10, free, c. 10–65 mm long, the filaments pubescent, eglandular; ovary glabrous and eglandular. Fruit a wingless, unarmed, coriaceous, glabrous, eglandular, oblong-elliptic, or linear pod, with a marcescent style forming an acute apex, c. 34–120 × 7–26 mm, explosively dehiscent, with twisting valves, 3–7-seeded. Seeds laterally compressed, obovate, up to 10 mm in diameter.

Caesalpinia, as re-circumscribed here, is reduced to around nine species (a detailed taxonomic revision is needed to properly delimit species), and is now restricted to the Neotropics (apart from the pantropically cultivated C. pulcherrima). All the Old World species previously included in Caesalpinia s.s. sensu Lewis (2005) are here transferred to other genera. One species (C. cassioides) occurs in the northern Andes from Peru to Colombia, one (C. pulcherrima) is likely native in Guatemala and the state of Sonora in Mexico), two occur in the Caribbean (one, C. nipensis, is endemic to Cuba, the other widely distributed and possibly divisible into six separate species, all of which are listed below). Caesalpinia pulcherrima is a widely cultivated ornamental throughout the tropics. It includes red, orange, pink, and pure yellow-flowered forms and cultivated specimens are usually unarmed and lack bristles (unlike wild specimens which are armed and bristly).

Seasonally dry tropical forests, coastal thicket, bushland and thorn scrub, dry plains and riparian woodland, on soils derived from limestone or sandstone.

Named by Linnaeus for Andrea Cesalpino (1519–1603), Italian naturalist, botanical collector, systematist and philosopher, physician to Pope Clement VIII, professor of medicine and botany in Pisa and Rome.

Britton and Rose (1930); Macbride (1943: 191, 194–195); Ulibarri (1996); Barreto Valdés (2013).

Figure 10. 

Caesalpinia cassioides Willd. A median leaflet B, C median leaflets (to show variation) D inflorescence E, F stem armature G flower H calyx opened out I calyx margin J median petal K upper lateral petal, L stamen M gynoecium N stigma O leaf and immature fruits P single immature fruit. A, D, E, Q from Mayolo 325 B, C, R from Silverstone-Sopkin 2004 F from Sandeman 4613 G–P from Silverstone-Sopkin 5139. Drawn by Sue Wickison.

Figure 11. 

Caesalpinia bahamensis Lam. A inflorescence D fruits (G. P. Lewis, Cuba, Lewis 1853 (K)). Caesalpinia nipensis Urb. B flowers E fruits (G. P. Lewis, Cuba, Lewis 1838 (K)). Caesalpinia cassioides Willd. C inflorescence (C. E. Hughes, Ancash, Peru, Hughes et al. 2228 (K)). Caesalpinia pulcherrima L. (Sw.) F inflorescence (C. E. Hughes, Sonora, Mexico, unvouchered); Denisophytum pauciflorum (Griseb.) E. Gagnon & G. P. Lewis G flower and leaves (G. P. Lewis, Cuba, Lewis 1854 (K)) H branch with spine-tipped woody protuberances (B. Torke, Cuba, Torke et al. 1424 (NY)). Denisophytum madagascariense R. Vig. I flowers and fruits (G. P. Lewis, Madagascar, Lewis et al. 2158 (K)). Gelrebia trothae (Harms) E. Gagnon & G. P. Lewis J inflorescence (P.J. Cribb, Tanzania, unvouchered).

Denisophytum is closely related to Tara (Fig. 3), but differs in having flowers with a lower cucullate sepal with an entire margin (vs. a lower cucullate sepal with a pectinate margin), and dehiscent, coriaceous, laterally compressed pods (except for D. madagascariense which has inflated fruits) (vs. indehiscent, somewhat fleshy, coriaceous pods that are slightly turgid). Morphologically, species of Denisophytum are most likely to be confused with those of Caesalpinia s.s., but no reliable diagnostic characters have been found to differentiate these two genera. The corolla of Denisophytum species is consistently yellow and the flowers are bee pollinated, whereas Caesalpinia s.s. species display a wide range of flower colour (yellow, orange, red, green and white) and pollination syndromes (chiropterophily, ornitophily, psychophily and mellitophily).

Denisophytum madagascariense R. Vig.

Shrubs to small trees, 0.5–2 (–5) m tall, armed with straight or curved, deflexed prickles, scattered along shoots and also in pairs at the petiole base (except D. madagascariense which is unarmed); young twigs glabrous to pubescent, eglandular. Stipules either minute or foliaceous and conspicuous, caducous (persistent in D. stuckertii). Leaves alternate, bipinnate, ending with a pair of pinnae; petiole and rachis glabrous and eglandular, with membranous or spinulose stipels at the insertions of pinnae on the leaf rachis, occasionally also at the insertion of the leaflets on the pinnae; pinnae opposite, in 1–6 pairs per leaf; leaflets opposite, in 2–10 (–11) pairs per pinna, elliptic, obovate to orbicular, with a rounded, acuminate or emarginate apex, c. 2–25 × 3–12 mm, leaflet blades glabrous to pubescent, eglandular. Inflorescence a terminal or axillary raceme; bracts caducous (acuminate and filiform in D. stuckertii). Flowers bisexual, zygomorphic; calyx a short hypanthium with 5 sepals, c. 4–10 mm long, eglandular, glabrous to finely pubescent, lower sepal cucullate and covering the other 4 sepals in bud, all sepals caducous, leaving a persistent free hypanthium ring on the pedicel as the fruit develops; petals 5, free, yellow, the median petal sometimes with red markings on the inner face of the blade, c. 5–10 mm long, obovate, petal claw almost absent (present in D. madagascariense); stamens 10, free, filaments pubescent and eglandular (8–11 mm long in D. madagascariense), anthers dorsifixed, glabrous to pubescent; ovary glabrous. Fruits coriaceous, oblong-elliptic, laterally compressed (but inflated in D. madagascariense), glabrous, eglandular pods with a tapering, sharp beak, 18–49 × 5–15 mm, elastically dehiscent, with twisting valves. Seeds ovoid, laterally compressed.

Denisophytum comprises nine taxa in eight species, found across North America, South America and Africa, including Madagascar, a classical highly disjunct trans-continental distribution typical of lineages occupying the succulent biome sensu Schrire et al. (2005). Three species are distributed in Mexico, Florida, and the Caribbean, one species is endemic to Paraguay and Argentina, one is endemic to northern Madagascar, and the other three occur in northern Kenya, Somalia and Arabia. An evaluation of species limits is needed in this group.

Low deciduous seasonally dry tropical woodland or scrubland, also in open pineland or coastal plains and foothills. Species in Madagascar and Africa grow in limestone soils.

There is no indication of the etymology of Denisophytum in the posthumous publication of the generic name. Nevertheless, it is quite likely that the author, René Viguier, had intended to honour his friend and collaborator, Marcel Denis, a botanist with expertise in the genus Euphorbia in Madagascar. Sadly, M. Denis passed away prematurely at the age of 33 in 1929 (Allorge and Allorge 1930).

Britton and Rose (1930); Burkart (1936: 84–86); Viguier (1949); Roti-Michelozzi (1957); Brenan (1967); Capuron (1967); Thulin (1983: 16–18; 1993: 344–347); Ulibarri (1996); Du Puy and Rabevohitra (2002); Barreto Valdés (2013).

Figure 12. 

Denisophytum stuckertii (Hassl.) E. Gagnon & G. P. Lewis. A foliage and inflorescences B median leaflet undersurface C stipule D leaf rachis spines E bract F calyx opened out G median petal H lateral petal I stamen J gynoecium K stigma L developing ovary M infructescence, N single fruit valve after dehiscence. A, B, D–K from Renvoize et al. 3538 C, M from Venturi 7697 L from Ruiz et al. 10488c N from Aguilar 241. Drawn by Eleanor Catherine.

Caesalpinia bessac Chiov., Flora Somala 1: 156. 1929.

SOMALIA, Uebi, Aug 1891, Robecchi-Bricchetti 622 (FI).

Denisophytum bessac is based on depauperate material and is of dubious status (Thulin, 1993).

Caesalpinia buchii Urb., Symb. Antill. 7(4): 510. 1913.

HAITI, “inter Gonaïves et Grosmorne ad Perou”, Buch 322 (holotype presumed at B†).

Caesalpinia erianthera Chiov., Fl. Somala 1: 155. 1929.

SOMALIA, from Obbia to Wuarandi, Aug 1891, Robecchi-Bricchetti 534 (syntype FI, fragments K!); and Boscaglia between Attod and Doldobscio, Apr 1924, Puccioni & Stefanini 450 (syntype FI).

Caesalpinia erianthera var. pubescens Brenan, Kew Bull. 17(2): 203. 1963.

KENYA, Northern Frontier Province, Banessa-Ramu, 23 May 1952, Gillett 13274 (holotype K!; isotype EA).

MADAGASCAR, Loky R. basin, Perrier de la Bâthie 4147 (holotype P).

Caesalpinia antsiranensis Capuron, Adansonia, sér. 2, 7: 203. 1967.

Type. MADAGASCAR, NE of Diego Suarez [Antsiranana], Orangea, Capuron 22990-SF (holotype P).

Libidibia pauciflora Griseb., Cat. Pl. Cub.: 78. 1866, (as “Lebidibia”).

Poinciana pauciflora (Griseb.) Small, Fl. SE United States: 59. 1903.

Caesalpinia pauciflora (Griseb.) C. Wright ex Sauvalle, Anal. Acad. Cienc. Med. Habana 5: 404. 1868 [1869].

Type. CUBA or. et occ., Wright 2361 (holotype ?GOET, n.v., isotype K!).

Caesalpinia rosei Urb., Repert. Sp. Nov. Regni Veg. 15: 314. 1918.

DOMINICAN REPUBLIC (Santo Domingo) prope Azua, Rose, Fitch & Russell 3861 (holotype US, photo K!).

Caesalpinia sessilifolia S. Watson, Proc. Amer. Acad. Arts and Sci. 21: 450 (1886).

Poinciana sessilifolia (S. Watson) Rose, in Contrib. U. S. Nat. Herb. 13(9): 303 (1911).

MEXICO, Bolson de Mapimi, 10 May 1847, Gregg s.n. (syntype NY); Mexico, Coahuila, on hills and mesas about Jumulco, May 1885, Pringle 202 (syntypes BR, CAS, CORD!, E, F, GH, GOET, JE, K!, MO, PH, SI!, US).

Caesalpinia stuckertii Hassl., in Repert. Sp. Nov. Reg. Veg. 12: 201 (1913).

ARGENTINA, Prov. Tucuman, Dept. Bunyacu: prope Cañada Alegre, 5 Jan 1900, Stuckert 21276 (? holotype SI).

Caesalpinia herzogii Harms, in Meded. Rijks-Herb. 27: 38 (1915).

Type. ARGENTINA, Gran Chaco: near Camoteras, Nov 1910, Herzog 1077 (? holotype L).

Caesalpinia stuckertii var. robusta Hassl., in Repert. Sp. Nov. Reg. Veg.12: 202. 1913.

Type. ARGENTINA, Prov. Tucuman, Depto. Bunyacu: Cañada Alegre, 31 Dec 1908, Stuckert 19726 (? holotype SI).

Tara differs from the closely related Coulteria in having racemose or paniculate inflorescences of bisexual flowers (vs. racemose inflorescences of unisexual flowers), indehiscent, laterally compressed, oblong, straight, slightly turgid and somewhat fleshy, coriaceous, sessile pods (vs. chartaceous to papyraceous, laterally-compressed, oblong to elliptic, occasionally suborbicular, pods, with a stipe ca. 4–13 mm long), and ellipsoid (vs. ovate-orbicular to sub-quadrate, compressed) seeds.

Tara tinctoria Molina ≡ Tara spinosa (Molina) Britton & Rose

Shrubs or trees, 3–5 (– 8) m tall, armed with deflexed prickles on the shoots; twigs glabrous to puberulent. Stipules not seen. Leaves alternate, bipinnate, ending with a pair of pinnae, sometimes armed with prickles at the base of the pinnae and leaflets; pinnae in 2–5 opposite pairs; leaflets opposite, in 1–8 pairs per pinna, obovate, broadly elliptic to oblong-elliptic, apex rounded, obtuse, to slightly emarginate, base equal or asymmetrical, rounded to cuneate, 10–46 × 7–35 mm, eglandular, glabrous or pubescent on lower surface; primary vein centric, secondary venation reticulate. Inflorescences in terminal or axillary racemes or panicles, rachis c. 5–30 cm long, glabrous or puberulous, eglandular, unarmed; bracts minute, usually under 3 mm long, with a long acuminate tip, caducous. Flowers bisexual, zygomorphic; calyx a hypanthium with five sepals that are 6–9 mm long, eglandular, glabrous to puberulous, lower sepal cucullate covering the other 4 sepals in bud, with a pectinate, fimbriate or entire margin, sepals caducous, but the hypanthium persisting as a calyx ring around the pedicel as the pod matures; petals 5, free, yellow, the median petal with red markings, c. 10 mm long; stamens 10, free, the filaments pubescent, eglandular. Fruit an indehiscent, straight, oblong, laterally compressed, slightly turgid and somewhat fleshy, coriaceous pod, 4–15 × 1.2–4 cm, eglandular, often puberulent when young, glabrescent. Seeds ellipsoid, c. 8–10 mm diameter, brown, shiny.

A genus of three species, one in South America (T. spinosa thought to be native to Peru and Ecuador), one in Mexico (T. cacalaco) and one in Mexico, Guatemala, Nicaragua and extending into the Caribbean (T. vesicaria). Tara spinosa is also widely cultivated across the tropics and subtropics (including in the Canary Islands) as a source of tannins and occasionally as an ornamental.

Seasonally dry tropical forest to semi-arid thorn scrub.

Derived from the vernacular name ‘tara’ in Peru, Bolivia and Chile.

Based on Gagnon et al. (2013), Molinari-Novoa and Sánchez Ocharan (2016) transfered C. cacalaco and C. vesicaria to the genus Tara, but did not emend the description of the genus, which we provide above.

Britton and Rose (1930); Sprague (1931); Macbride (1943, as Caesalpinia spinosa, 195-196); Ulibarri (1996); Barreto Valdés (2013); Molinari-Novoa and Sánchez Ocharan (2016).

Figure 13. 

Tara spinosa (Molina) Britton & Rose. A habit B leaflet undersurface, C section of young stem D flower E calyx opened out F median petal G upper lateral petal H lower lateral petal I stamen J gynoecium K stigma L fruit M seed. A–K from Lewis 1416 L, M from Filskov et al. 37341. Drawn by Eleanor Catherine.

Figure 14. 

Caesalpinia (Coulteria) velutina Britton & Rose. A inflorescence (G. P. Lewis, Guatemala, Lewis et al. 1713 (K)) B fruits (C. E. Hughes, Guatemala, Lewis et al. 1714 (K)). Tara vesicaria (L.) Molinari, Sánchez Och. & Mayta C habit (C. E. Hughes, Tecolostote, Nicaragua, Hughes 1376 (FHO)) H flower (C. E. Hughes, Rivas, Nicaragua, J. A. Hawkins 11 (FHO). Tara spinosa (Molina) Britton & Rose D inflorescence (E. Gagnon, Ancash, Peru, Hughes et al. 3043 (MT)) I unripe fruits (C. E. Hughes, Cajamarca, Peru, Hughes 1996 (FHO)). Tara cacalaco (Humb. & Bonpl.) Molinari & Sánchez Och. E flowers (C. E. Hughes, Puebla, Mexico, Hughes et al. 2169 (FHO)) F unripe fruits (G. P. Lewis, Mexico, MacQueen 488 (K)) G bark (C. E. Hughes, Puebla, Mexico, Hughes et al. 2073 (FHO)).

Coulteria differs from Tara by its racemose inflorescences of unisexual flowers (vs. inflorescences of racemes and panicles with bisexual flowers), chartaceous to papyraceous, laterally-compressed, oblong to elliptic (occasionally suborbicular) stipitate pods, subtended by a 4–13 mm long stipe (vs. indehiscent, laterally compressed but slightly turgid and somewhat fleshy, coriaceous, straight, oblong, sessile pods), and compressed, ovate-orbicular to sub-quadrate, compressed (vs. ellipsoid) seeds.

No type designated in the original publication, nor since. Type designated here: Coulteria mollis Kunth.

Trees or shrubs, 3–20 m tall, unarmed; young twigs with a dense velvety-bronze pubescence, glabrescent. Stipules not seen. Leaves alternate, bipinnate, ending in a pair of pinnae; petiole and rachis glabrous or densely velutinous; pinnae in 2–6 pairs; leaflets in (2–) 4–12 (– 14) pairs per pinna, 0.6–8 cm long, elliptic, oblong to ovate, apex obtuse to acute, base narrow, rounded or obtuse, eglandular, glabrous to velvety pubescent; main vein centric, secondary veins brochidodromous. Inflorescence racemose, axillary or terminal, 5–16 (– 25) cm long; bracts minute, with an acute tip, pubescent, caducous. Flowers unisexual, male and female flowers on separate trees, zygomorphic; calyx comprising a hypanthium with 5 sepals, 8–10 mm long, velvety-pubescent, lower sepal cucullate, glandular-pectinate, covering the other 4 sepals in bud; petals 5, yellow, free; male flowers with 10 free stamens, filaments pubescent, eglandular. Fruit chartaceous to papyraceous, laterally-compressed, oblong to elliptic (occasionally suborbicular), indehiscent (or sometimes opening along one suture), wingless, 3–15 × 2–4 cm, with a 4–13 mm long stipe, pendulous, often persisting to next flowering season, eglandular, glabrous to densely velutinous, 1–6-seeded. Seeds ovate orbicular or sub-quadrate, compressed.

A genus of approximately seven species in Mexico and Central America, one species extending to Cuba, Jamaica and Curaçao, one to Venezuela (including Isla Margarita) and Colombia.

Seasonally dry tropical forest, deciduous woodland and dry thorn scrub, some species occurring on limestone.

Named by Kunth for the Irish botanist Thomas Coulter (1793–1846) who collected in central Mexico (1825–1834) and was curator of the herbarium at Trinity College, Dublin, Ireland.

A revision of the genus has been submitted by S. Sotuyo, J. L. Contreras, E. Gagnon, and G. P. Lewis. The list of species names presented here simply includes all names associated with the genus Coulteria and will be reduced in the forthcoming taxonomic account.

Britton and Rose (1930: 320–322); Ulibarri (1996); Zamora Villalobos (2010); Sotuyo et al. (submitted)

Figure 15. 

Caesalpinia (Coulteria) velutina Britton & Rose. A portion of leaf B detail of bark C inflorescence D flower E calyx opened out F detail of calyx lobe G median petal H upper lateral petal I lower lateral petal J stamen K fruit L seed M seedling. A, K from Lewis and Hughes 1714 B–J, M from Lewis and Hughes 1713. Drawn by Eleanor Catherine.

Gelrebia is morphologically similar to Caesalpinia s. s. but the two genera differ somewhat in habit, with Gelrebia species being erect to scrambling shrubs (vs. erect shrubs or small trees), in having dark pinkish mauve to light pinkish-white flowers (vs. flowers that are variable in colour, from yellow, white, red and orange to green), and coriaceous, broadly oblong-ovoid to obliquely pyriform pods, with a large, oblique, rounded base (vs. coriaceous, oblong-elliptic to linear pods, with an oblique cuneate base).

Gelrebia rubra (Engl.) E. Gagnon & G. P. Lewis ≡ Hoffmannseggia rubra Engl.: Caesalpinia rubra (Engl.) Brenan

Erect to scambling shrubs, 0.3–5 m tall, armed with scattered, straight or curved, deflexed prickles (these 7–20 mm long); stems puberulous to pubescent when young, glabrescent. Stipules not seen. Leaves alternate, bipinnate, ending in a pair of pinnae; pinnae opposite, in 1–17 pairs; leaflets opposite (except in G. glandulosopedicellata), in 1–33 pairs per pinna, narrowly oblong or oblong-elliptic, 3–11 × 2–5 mm, apex rounded to emarginate, sometimes mucronate, glabrous or sparsely pubescent, lower surface of the blades with numerous subepidermal glands or translucent dots (best seen with a × 10 hand lens or microscope). Inflorescence a terminal or axillary raceme, c. (1–) 2–19 (– 25) cm long, unarmed; bracts broadly ovate to suborbicular, apex aristate, 3–10 mm long, caducous. Flowers bisexual, zygomorphic; calyx comprising a short hypanthium with 5 sepals, c. 5–13 mm long, eglandular, glabrous to finely pubescent, lower sepal strongly cucullate (occasionally with a beaked apex), covering the other 4 sepals in bud before anthesis, all sepals caducous, but hypanthium persisting as a free ring around the pedicel as the pod matures; petals 5, free, dark pinkish mauve to light pinkish-white, c. 7–24 × 5–15 mm, eglandular; stamens 10, free, filaments 8–20 mm long, pubescent and eglandular; ovary glabrous. Fruit a coriaceous, broadly oblong-ovoid to obliquely pyriform pod, apex acute, with a large, oblique, rounded base, c. 15–40 × 12–23 mm, dehiscent along both sutures, glabrous to minutely pubescent, eglandular. Seeds obovoid, laterally compressed.

A genus of nine taxa in eight species, restricted to Africa, in Namibia, Botswana, South Africa, Northern Kenya, Ethiopia, and Somalia. One species also found in the Democratic Republic of the Congo (Zaire, Katanga).

Deciduous bushland, dry woodlands, on rocky ridges, often along dry river beds, or on sandy valley floors. One species also found in degraded savanna, close to termite mounds.

Gelreb or gelrib is the Somali name for Gelrebia trothae subsp. erlangeri (field labels of Dale K724 (“gelrib”) and of Gillett 13223 (“gelreb”) from Kenya), meaning ‘camel trap’ and clearly alluding to the robust deflexed prickles characteristic of the species, and indeed the genus as a whole, which can hinder the passage of camels.

Wilczek (1951); Roti-Michelozzi (1957); Brenan (1963, 1967); Ross (1977: 122–130); Thulin (1980, 1983: 16–18; 1993: 344–347); Germishuizen (1991); Roux (2003); Curtis and Mannheimer (2005: 226–228); Brummitt et al. (2007).

Figure 16. 

Gelrebia trothae E. Gagnon & G. P. Lewis subsp. trothae. A part of branch showing inflorescence with flowers and fruits B portion of leaflet margin, lower surface C longitudinal section of flower D median petal inner surface E lateral petal inner surface F stamen G anther H ovary with part of wall removed to expose ovules I fruit valve after dehiscence J seed. Gelrebia trothae subsp. erlangeri (Harms) E. Gagnon & G. P. Lewis K part of inflorescence L fruit. A–H from Milne-Redhead & Taylor 11177 I, J from Ward U27 K from Gillett 13223 L from Hemming 478. Drawn by L. M. Ripley, originally published in F.T.E.A., Leguminosae subfamily Caesalpinioideae, page 34, fig. 5 (1967).

Caesalpinia bracteata Germish., Bothalia 21 (2): 153. 1991.

[South Africa, Cape Province]: “2819 (Ariamsvlei): Kenhardt District, on farm Skroef, near hot spring (Warmbad Noord) on Orange River (-DA)”, 29 Sep 1987, Van Hoepen 1941 (holotype PRE).

Caesalpinia dauensis Thulin, Kew Bull. 34(4): 819. 1980.

KENYA, 30 km on the Ramu-Malka road, c. 4°04'N, 40°59'E, 8 May 1978, Gilbert & Thulin 1583 (holotype UPS; isotypes BR, EA, K!).

Caesalpinia glandulosopedicellata R. Wilczek, Bull. Jard. Bot. Brux. 21: 83. 1951.

“Congo Belge”, district du Haut-Katanga: environs de Niemba, Schmitz 1595.

Caesalpinia merxmuellerana A. Schreib., Mitt. Bot. St. Munchen 16, Beih., Die Gattung Caesalpinia in Südwestafrica, 64. 1980.

SOUTH WEST AFRICA, Dist. Lüderitz-Süd, Farm Uitsig, Wendt in herb. W. Giess 14713 (holotype M; isotypes K!, PRE, WIND).

Caesalpinia oligophylla Harms, Engl., Bot. Jahrb. Syst. 33: 160. 1902.

ETHIOPIA, “Arussi Galla”, Apr 1901, Ellenbeck 2038 (holotype B †); SOMALIA, rive dello Scebelia Bulo Burti, 25 Feb 1924, Puccioni & Stefanini 134 (neotype FI, designated by G. Roti-Michelozzi in Webbia 13: 207. 1957).

Caesalpinia rostrata N. E. Br., Hooker's Icon. Pl., 28: t. 2702. 1901.

SOUTH AFRICA, from cultivation in Durban Botanic Garden, raised from seed obtained from “Delagoa Bay”, Maputo (Lourenço Marques), Wood 7943 (holotype K!; isotypes BOL, NH, PRE).

Hoffmannseggia rubra Engl., Bot. Jahrb. Syst. 10: 25. 1889. Caesalpinia rubra (Engl.) Brenan, Kew Bull. 17(2): 202. 1963.

NAMIBIA, Karibib Dist., Usakos, Marloth 1432 (holotype ?B; isotypes BOL, PRE).

Caesalpinia trothae Harms, Engl., Bot. Jahrb. Syst., 26: 277. 1899, as “trothaei”.

TANZANIA, ?Dodoma District, Ugogo, Chumo Pass, Jan. 1897, von Trotha 186 (holotype B †).

Caesalpinia erlangeri Harms, Engl., Bot. Jahrb. Syst. 33: 160. 1902.

Caesalpinia trothae subsp. erlangeri (Harms) Brenan, Kew Bull. 17(2): 20. 1963.

ETHIOPIA, Galla Sidama, Borana, Tarro Gumbi, Ellenbeck 2071 (holotype B †). Somalia, Dolo, sul Daua, 6 May 1893, Riva 1104 (neotype FI, designated by G. Roti-Michelozzi in Webbia 13: 209, 1957).

Hultholia is closely related and morphologically similar to Guilandina. While both genera form armed lianas, Hultholia differs in having stems with dome-shaped glands intermixed with dense slender, patent, needle-like prickles (vs. stems eglandular and with strongly recurved, robust prickles in Guilandina); both genera have sharp recurved prickles on the leaf and pinnae rachises. Hultholia has bisexual flowers (vs. unisexual flowers on separate female and male racemes in Guilandina), a zygomorphic corolla, with petals extending beyond the sepals, and the median (standard) petal smaller than the other four (vs. a sub-actinomorphic to zygomorphic corolla, with petals only slightly extending beyond the sepals in Guilandina), unarmed, obovoid, falcate, pubescent, vesicular pods (vs. oblong-elliptic, coriaceous, eglandular, inflated pods, usually armed with 5–10 mm long, slender spinescent bristles), and sub-globose, oblong, grey, ca. 10 × 7 mm, smooth seeds (vs. obovoid to globular c. 20 mm in diameter, grey, pale to dark brown or orange seeds, with parallel fracture lines concentric with the small apical hilum).

Hultholia mimosoides (Lam.) E. Gagnon & G. P. Lewis ≡ Caesalpinia mimosoides Lam.

Climbing woody shrub; branches densely armed with short, robust, needle-like trichomes; young stems pubescent, with rust-coloured, hyaline hairs and dome-shaped glands, topped with a few hairs. Stipules subulate, 7–15 mm long, pubescent, caducous. Leaves alternate, bipinnate, without a single terminal pinna, 22–40 cm long; pinnae opposite, in 10–30 pairs per leaf, about 3–5 cm long, pubescent, with a pair of deflexed prickles at the insertion of the pinnae on the leaf rachis, and at the insertion of leaflets on the pinnae rachises; leaflets opposite, in 7–20 pairs per pinna, oblong, asymmetric at base, c. 9 × 4 mm, glabrous, eglandular. Inflorescences terminal or leaf-opposed, lax racemes, with 50 or more flowers, 20–40 cm long; rachis and pedicels armed with needle-like, robust trichomes, pubescent and covered with domed, hair-tipped glands. Flowers bisexual, zygomorphic; calyx comprising a hypanthium with 5 sepals 13–16 × 6 mm; hypanthium and sepals pubescent and glandular, the sepal margins sometimes with small stipitate glands, < 1 mm long; petals 5, free, bright yellow, dark glands present on the blade, median (standard) petal c. 8 mm wide and smaller than the 4 lateral petals, that are c. 1.7 × 1.3 cm; stamens 10, free, filaments 1.8 cm long, pubescent at least on the lower ½; ovary densely pubescent, and with glandular dots (often obscured by the dense pubescence). Fruit an obovoid, falcate, vesicular, unarmed, dehiscent pod, sparsely pubescent, particularly along the margin, and with a few obscure stellate hairs, and covered in gland dots, 5–6 × 2.5–3 cm, 1–3-seeded. Seeds sub-globose, oblong, 10 × 7 mm, grey.

The single species is distributed across Asia, in China (Yunnan), Bangladesh, India, Laos, Myanmar (Burma), Thailand and Vietnam.

In secondary thickets and clearings, often on roadsides, up to 1500 m elevation. More information on the ecology of this genus is needed.

The name Hultholia honours the Cambodian botanist Dr. Sovanmoly Hul Thol (born 1946), whose doctoral thesis, “Contribution à la révision de quelques genres de Caesalpiniaceae, representés en Asie” (1976), is an important revision of the Asian species and genera of the Caesalpinia group, and particularly the genus Pterolobium. Dr. Hul Thol retired from the Museum National d’Histoire Naturelle, Paris in 2014, but continues as an honorary researcher. She is a specialist on the flora of Cambodia and South East Asia, directed the publication of multiple volumes of the Flora of Cambodia, Laos and Vietnam from 1995, and is one of the co-founders of the National Herbarium of Cambodia, Royal University of Phnom Penh.

Although Hultholia mimosoides is not known to be cultivated, the young, pungent, flowering shoots are sold as a vegetable in markets in Vientiane (Laos) (Vidal and Hul Thol 1976).

Vidal and Hul Thol (1976); Chen et al. (2010a: 42–43).

Figure 17. 

Hultholia mimosoides (Lam.) E. Gagnon & G. P. Lewis. A habit, including foliage and inflorescences B stem armature detail C bud showing cucullate lower lobe of calyx D calyx lobes outer surface E calyx cucullate lower lobe side view, F median petal inner surface G median petal side view H upper lateral petal inner surface I lower lateral petal inner surface J stamens K anthers dorsal and ventral views L gynoecium M stigma detail N fruit O seed. A–K from Clark 237 L, M from Beusekom & Geesink 4706 N, O from Bunchuai 1342. Drawn by Juliet Williamson.

Figure 18. 

Hultholia mimosoides (Lam.) E. Gagnon & G. P. Lewis. A young leaves and inflorescence in bud (J. Jose, Wikicommons (https://commons.wikimedia.org/wiki/File:Caesalpinia_mimosoides_2_at_Kudayathoor.jpg), Kerala, India, unvouchered) B flower (R. Clark, Thailand, Clark et al. 237 (K)) C flowers D immature fruits E mature fruit F habit G open fruit with seeds (V. R. Vinayaraj, Wikicommons (https://commons.wikimedia.org/wiki/Category:Caesalpinia_mimosoides, the basionym of Hultholia mimosoides), India, unvouchered).

Caesalpinia mimosoides Lam., Encycl. Méth., Bot. 1(2): 462 (1785).

Biancaea mimosoides (Lam.) Tod., Hort. Bot. Panorm. 1(1): 3 (1875).

Specimen originally from Malabar, sent to Lamarck by Sonnerat (P: Herb. Lamarck, fide Vidal and Hul Thol. 1976).

Guilandina bonduc L.

Lianas, woody climbers, scrambling or trailing shrubs, often forming dense tangled clumps, densely armed with recurved prickles on branches and shoots, as well as in pairs at leaf bases (except Caesalpinia murifructa and closely related species in the Caribbean which are unarmed). Stipules foliaceous to subulate, sub-persistent or caducous. Leaves bipinnate, ending with a pair of pinnae, prickles present in pairs at the insertion of pinnae and scattered on the leaf rachis, and at the insertion of leaflets on the pinnae rachises; leaflets oblong, apex obtuse and mucronulate to acuminate, base rounded. Inflorescences supra-axillary or terminal racemes, 30–60 cm long; bracts narrow, lanceolate, aristulate, 1 mm long, to conspicuous and exceeding floral buds, caducous. Flowers unisexual, segregated on separate male and female racemes, the female flowers cryptically bisexual with 10 fully formed stamens, but these produce no pollen; male flowers with a highly reduced, non-functional pistil, zygomorphic to sub-actinomorphic; calyx with a hypanthium and 5 almost equal sepals, these valvate in bud, the lower sepal slightly cucullate, the hypanthium and sepals caducous, leaving no persistent calyx ring, eglandular, without spines (except Madagascan Caesalpinia delphinensis in which the calyx is armed with slender prickles); petals 5, free, yellow, barely exceeding the sepals; stamens 10, free, pubescent near the filament base; ovary usually covered in bristly trichomes, except in a few species, including Caesalpinia solomonensis and Caesalpinia murifructa. Fruits oblong-elliptic, inflated pods, usually armed with 5–10 mm long spinescent bristles, apex terminating in a beak, base acute, 1–4-seeded. Seeds obovoid to globular, c. 2 cm in diameter, smooth, grey, pale to dark brown, or orange, with parallel fracture lines concentric with the small apical hilum.

This pantropical genus lacks a recent global taxonomic account and there are doubts about the number of species, with previous estimates ranging from seven to as many as 19. Species occur from as far north as Japan, south to South Africa, with three species in the Caribbean, one in China, India, Myanmar (Burma), Indo China, Hong Kong and Taiwan, one endemic to Madagascar, one in Australia, and two widespread across the Old and New World tropics.

Coastal thickets on sand, in secondary forest, and lowland rain forest, occasionally on limestone.

Named by Linnaeus for Melchior Wieland (1515–1589), Prussian naturalist, traveller and scholar from Königsberg, who settled in Italy and italianised his name to ‘Guilandini’, or Guilandinus in Latin; he was sent to the Levant, Asia and Africa (1559–1560), was captured by pirates and finally ransomed by Gabriele Falloppio.

Pending a complete taxonomic revision, the list of 19 names presented below provides a guide to potential species content in Guilandina, but includes no synonymy and no information on types, nor any new nomenclatural combinations for the five species of Caesalpinia that as yet have no published name in Guilandina.

Britton and Rose (1930: 336–341); Wilczek (1951); Brenan (1967); Gillis and Proctor (1974); Hattink (1974); Vidal and Hul Thol (1976); Du Puy and Rabevohitra (2002: 46–48); Chen et al. (2010a).

Figure 19. 

Guilandina ciliata Bergius ex Wikstrom. A foliage B leaflet undersurface C prickle enlarged to show indumentum D inflorescence and portion of leaf; E flower F, G median petal H upper lateral petal (outer surface) I lower lateral petal (inner surface) J stamens K stamen L fruit M, N seeds. A–C from Ekman 5413 D–K from Curtiss 143 L–N from Pannell 179. Drawn by Pat Halliday.

Figure 20. 

Moullava spicata (Dalzell) Nicolson. A inflorescences B fruit (P. Awale, Flowers of India (http://www.flowersofindia.net/), Maharashtra, India, unvouchered) C flowers (M. Sanjappa, India, unvouchered). Guilandina bonduc L. D young fruits (F. Starr and K. Starr, Starr Environmental (http://www.starrenvironmental.com/images/species/?q=Caesalpinia+bonduc), Florida, USA, unvouchered) E fruits with mature seeds (G. P. Lewis, Madagascar, Du Puy et al. M665 (K)) F inflorescence (M. Sanjappa, India, unvouchered). Biancaea decapetala (Roth) O. Deg. G fruits with seeds H fruit with thickened suture (C. E. Hughes, Ancash, Peru, Hughes et al. 2227 (FHO)) I inflorescence (E. Gagnon, Ancash, Peru, Hughes et al. 3055 (MT)). Biancaea godefroyana (Kuntze) Molinari, Mayta & Sánchez Och. J inflorescences and fruits (F. Xaver, Wikicommons (https://commons.wikimedia.org/wiki/File:Caesalpinia_godefroyana_1.jpg), Cambodia, unvouchered).

Moullava is related to Mezoneuron, but differs by its fleshy, oblong-elliptic, indehiscent, sub-torulose, wingless pods, with thickened sutures (vs. laterally compressed, chartaceous, coriaceous or ligneous, indehiscent pods, with a longitudinal wing along the upper suture), and by its subglobular (vs. compressed) seeds.

“H.M. 6 t. 6” (= Rheede`s Hortus Malabaricus 6, plate 6, 1686) = Moullava spicata.

Lianas and scrambling shrubs, armed with deflexed prickles on shoots. Stipules not seen. Leaves alternate, bipinnate, ending with a pair of pinnae, 12–40 cm long, glabrous to pubescent-tomentose, with a pair of prickles at the insertion of each pinna; pinnae opposite, in 7–20 pairs; leaflets in 5–40 opposite pairs per pinna, sessile, narrowly oblong to ovate-oblong, apex rounded to emarginate, sometimes mucronate, base asymmetrical to rounded, blades eglandular, glabrous to pubescent, 4–20 × 2–6 mm. Inflorescence an elongated terminal or axillary raceme, the flowers subsessile, pedicels, when present, 10–25 mm long, the racemes sometimes aggregated into panicles, 8–60 cm long, unarmed or with a few prickles at the base. Flowers bisexual, sub-actinormophic or zygomorphic; calyx comprising a hypanthium with 5 sepals, 6–12 × 2–4 mm, the lower sepal strongly cucullate, covering the other 4 sepals in bud, all sepals eglandular and glabrous; petals 5, free, yellow, the median and lateral petals sometimes streaked red, eglandular; stamens 10, free, barely exserted beyond the corolla, densely pubescent on lower half of filaments, 8–15 mm long; ovary glabrous or pubescent. Fruit fleshy, oblong-elliptic, unarmed, indehiscent, sub-torulose, with thickened sutures, the apex apiculate, 35–50 (–80) × 15–30 mm, drying black (immature fruits of M. spicata red-tomentose), exocarp and endocarp strongly adnate, glabrous, 1–4-seeded. Seeds sub-globular, 12–20 mm in diameter, olive-brown to black.

A genus of four species, three in south Asia: India, Nepal, Myanmar (Burma), Thailand, Laos, Cambodia, Sri Lanka, southern China (Yunnan and Hainan), and the Malay Peninsula and Archipelago, and one in Africa: Cameroun, Gabon, the Democratic Republic of Congo, Angola, Zambia (Kabompo Dist.), Uganda and Tanzania (Kigoma Dist.).

The Asian species are found in seasonally dry tropical semi-evergreen forest margins, secondary thickets, and on mountain slopes, up to 1200 m elevation. The African species occurs mostly in riverine habitats in lowland rainforests.

Derived from the vernacular name of Moullava spicata, “mulu” (Malayalam: spiny), a spiny climber.

Brenan (1963, 1967); Hattink (1974); Vidal and Hul Thol (1976); Nicolson (1980); Ansari (1990); Sanjappa (1992: 33); Brummitt et al. (2007, see both Moullava and Mezoneuron welwitschianum); Chen et al. (2010a).

Figure 21. 

Moullava spicata (Dalzell) Nicolson. A flowering branch B single pinna of bipinnate leaf C leaflet undersurface D leaflet undersurface detail E young stem F older stem G part inflorescence H calyx opened out I median petal J upper lateral petal K lower lateral petal L stamen M gynoecium N stigma O fruit P seed. A, G from photo by P. S. Green B–D, H–N from Cult. Foster Bot. Gard. F1901, specimen Hutchinson 2784 E from Critchett 11/79 F from Nana 5620 O, P from Meebold 8605. Drawn by Eleanor Catherine.

Caesalpinia digyna Rottl., Ges. Naturf. Freude Berlin Neue Schriften 4:198–200, pl. 3. 1803.

[S. INDIA] Marmelon (near Madras), 9 Oct 1799, Rottler s.n. (? B: Herb. Willdenow, K!).

Caesalpinia gracilis Miq., Fl. Ned. Ind. 1:110. 1855.

Type. INDIA, Roxburgh (n.v.).

Caesalpinia oleosperma Roxb., Hort. Bengal. 32. 1814.

Type. JAVA, Horsfield 138 (holotype K!; isotype BM).

Caesalpinia flavicans Grah., Cat.: 5825. 1832, nom. nud.

Caesalpinia spicata Dalzell, in Hooker’s J. Bot. Kew Gard. Misc. 3: 89 (1851).

Wagatea spicata Dalzell, in Hooker’s J. Bot. Kew Gard. Misc. 3: 89 (1851).

WESTERN INDIA, Bombay presidency.

Caesalpinia ferox Hohen., Pl. Ind. Or. Exs. No. 414, non Hassk.

Type. Not traced.

Caesalpinia digyna Graham, Cat. 60. 1839, non Rottl. 1803, nom. illeg.

Caesalpinia mimosoides Heyne & Wall, Numer. List n. 5837. 1831, nom. illeg., non Lam.1785.

Caesalpinia tortuosa Roxb., Fl. Ind. (ed. 1832) 2: 365. 1832.

Specimen originating from SUMATRA, cultivated in the Botanic Garden of Calcutta, “Hort. Calc. E. Sumatra”, Roxburgh s.n. (holotype: K!).

Caesalpinia acanthobotrya Miq., Fl. Ned. Ind. 1(Suppl.): 108 (1860) & 293 (1861).

Type. W. SUMATRA, prov. Priaman, 1855–60, Diepenhorst HB2240 (holotype U; isotype BO).

Caesalpinia microphylla Buch.-Ham ex Prain, in J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 66: 471. 1897, non Mart. ex G. Don, 1832.

Type. INDIA, Goyalpara, 6 Aug 1908, Wallich 5826 (K!).

Caesalpinia tortuosa var. grandifolia Craib, Fedde Repert. Spec. Nov. Reg. Veg. 12: 392. 1913.

Type. MYANMAR [Burma], Kowpok, Jan 1912, Meebold 17208 (K!).

Caesalpinia cinclidocarpa Miq., in Fl. Ned. Ind 1: 110 (1855).

Type. JAVA, as for Cinclidocarpus nitidus, non Caesalpinia nitida Hassk. (1844).

Cinclidocarpus nitidus Zoll. & Moritzi, in Naturr-Geneesk. Arch. Ned.-Indie 3: 82 (1846).

Type. JAVA, Zollinger 3462 (holotype L; isotypes A, BM, P).

Caesalpinia tortuosa Wall., Numer. List n. 5827 D. 1831, nom. nud.

Mezoneuron welwitschianum Oliv., Fl. Trop. Afr. 2: 261. 1871.

Caesalpinia welwitschiana (Oliv.) Brenan, Kew Bull. 17(2): 203. 1963.

ANGOLA, Cuanza Norte, Golungo Alto, Welwitsch 608 (holotype LISU; isotypes BM, K!).

Biancaea is closely related to Mezoneuron, differing principally in its fruit, a coriaceous, laterally compressed, wingless, dehiscent pod (except B. decapetala, which has somewhat inflated, boat-shaped pods, often with a narrow wing or ridge along the upper suture). In contrast, Mezoneuron has chartaceous, coriaceous or ligneous pods, which are also laterally compressed, but indehiscent, and with a wing along the upper suture. In addition, the ovary of Biancaea species always has a velvety indumentum (vs. glabrous to pubescent in Mezoneuron).

Biancaea scandens Tod. ≡ Biancaea decapetala (Roth) Deg.

Lianas, climbing or trailing shrubs (1–3 m), or small trees (2.5–10 m), armed with short, slightly recurved prickles, scattered along the branches; young shoots pubescent or glabrescent. Stipules lanceolate-oblong to broadly-ovate, sometimes amplexicaul at base, 3–4 mm to 4.5 cm long, caducous or sub-persistent to persistent. Leaves alternate (except in B. oppositifolia), bipinnate, ending with a pair of pinnae, rachis pubescent (glabrous in B. oppositifolia), armed with pairs of prickles at the base of each pinna, sometimes also scattered on the rachis; pinnae in 4–19 opposite to alternate pairs; leaflets opposite to alternate, in 5–20 pairs per pinna, blade membranous, eglandular, glabrous to pubescent, 10–35 × 4–15 mm (4–10 × 1.5–4.5 cm in B. oppositifolia), oblong-elliptic, apex acute, obtuse, rounded to emarginate, base asymmetric. Inflorescences erect, showy, terminal or axillary racemes or panicles; rachis eglandular, pubescent, unarmed or with a few scattered prickles, mainly near the base; bracts ovate-lanceolate, acuminate, 2–8 mm long, caducous. Flowers bisexual, zygomorphic; calyx with a short hypanthium and 5 sepals, the lower sepal cucullate and covering the other 4 in bud, sepals pubescent (except in B. sappan), caducous, but the hypanthium persisting as a calyx ring around the pedicel as fruits mature; petals 5, free, yellow to white, eglandular, the claws pubescent; the median petal smaller than the other 4, and inrolled towards the centre, lateral petals oblong, obovate to spathulate, 4–10 × 2–8 mm; stamens 10, filaments densely pubescent (most evident at the base), eglandular, 10–15 mm long; ovary densely velutinous. Fruit a coriaceous, glabrous, eglandular, oblong-elliptic to obovate, dehiscent, wingless, laterally compressed (but somewhat inflated and often with a narrow wing along the upper suture in B. decaptala), 4.5–10 × 2–4 cm, 2–8-seeded pod, usually much broader at the rounded to truncate apex, which terminates in a sharp beak. Seeds flat, elliptic, ovoid to orbicular, c. 2 cm in diameter, black or brown.

A genus of six species widespread across southern Asia, from India, to Myanmar (Burma), Thailand, Cambodia, Vietnam, south China, Japan, the Philippines, and the Malay Peninsula and Archipelago, one species endemic to Sabah (near Sandakan). Biancaea decapetala, native to Asia, has been widely introduced across the tropics as a hedge plant or ornamental and is considered to be invasive in South Africa and Hawaii.

Primary forest and forest margins, grasslands, scrub vegetation, riverine habitats, secondary thickets and clearings. From the coast to mountain slopes.

Unknown.

Based on the study of Gagnon et al. (2013), Molinari-Novoa et al. (2016) provided some, but not all, of the required nomenclatural transfers to the genus Biancaea. Furthermore, they did not emend the description of the genus, as provided here.

Hattink (1974); Vidal and Hul Thol (1976); Jansen (2005); Brummitt et al. (2007); Chen et al. (2010a); Molinari-Novoa et al. (2016).

Figure 22. 

Biancaea decapetala (Roth) O. Deg. A flowering branchlet and foliage B, C leaflets viewed from above and below, respectively D flower with parts separated, and centre of flower enlarged E calyx three views F lateral petal G median petal H stamen I anther J gynoecium K fruits L seed. A from Rutherford-Smith 11062 B, C from White 2478 D–J from Chase 4564 K, L from Myre 2528. Drawn by D. Erasmus, originally published in Flora Zambesiaca, vol. 3 part 2, page 182, figure 3.2.39 (2007).

Reichardia decapetala Roth, Nov. Pl. Sp. 212. 1821.

Caesalpinia decapetala (Roth) Alston, Handb. Fl. Ceylon 6: 89. 1931.

Type. INDIA, (fl.), Heyne s.n. (isotype K!).

Biancaea scandens Tod., in Nuov. Gen. Sp. Pl.: 22. 1860.

Type. “Cortivasi da lungo tempo nel Real Orto Botanico [di Palermo] in piena terra, col nome di Caesalpinia sepiaria”.

Caesalpinia benguetensis Elmer, in Leafl. Philipp. Bot. 1: 226 (1907).

Mezoneuron benguetense (Elmer) Elmer, in Leafl. Philipp Bot 1: 362 (1908).

Type. PHILIPPINES, Luzon, Benguet prov. Baguio, (fl. fr.), Mar 1907, Elmer 8720 (BO, K!, L, PHN).

Caesalpinia japonica Sieb. & Zucc., in Abh. Math.-Phys. Cl. Königl. Bayer Akad. Wiss. 4(2): 117. 1845.

Caesalpinia sepiaria var. japonica (Siebold & Zucc.) Gagnep., in Fl. Indo-Chine 2: 180. 1913.

Caesalpinia sepiaria var. japonica (Siebold & Zucc.) Makino, Ill. Fl. Nippon: 431. 1940.

Caesalpinia decapetala var. japonica (Siebold & Zucc.) H. Ohashi, Fl. E. Himalaya 3: 58. 1975.

Caesalpinia decapetala var. japonica (Siebold & Zucc.) Isely, Mem. New York Bot. Gard. 24(2): 193. 1975.

Type. JAPAN, Siebold & Zuccanini.

Caesalpinia ferox Hassk., Ind. Sem. Hort. Amst. 1841.

Biancaea ferox (Hassk.) Tod., Hort. Bot. Panorm. 1(1): 3. 1875.

Type. probably a living plant in Hort. Bog., fide Hattink (1974).

Caesalpinia sepiaria Roxb., Fl. Ind. 2: 360. 1832. Biancaea sepiaria (Roxb.) Tod., Hort. Bot. Panorm. 1(1): 3. 1875.

Type. INDIA, Roxburgh without number (isotypes: BM, K!, in Hb. Wallich 5834A).

Caesalpinia sepiaria Roxb. var. pubescens T. Tang. & F.T. Wang, Illust. Treat. Prin. Pl. China (Leguminosae): 96. 1955, without Latin description.

Caesalpinia sepiaria Roxb. var. pubescens T. Tang & F. T. Wang ex C. W. Chang, Flora Tsinlingensis 1(3): 444. 1981.

Caesalpinia decapetala (Roth) Alston var. pubescens P. C. Huang, Sylva Sinica 2: 1187. 1985, nom. illeg., without Latin description or type.

Caesalpinia decapetala var. pubescens (T. Tang & F. T. Wang ex C. W. Chang) X. Y. Zhu, in Legumes of China: 5. 2007.

Type. CHINA.

Caesalpinia godefroyana Kuntze, Rev. Gen. Pl. 1: 166. 1891.

VIETNAM (South), Cap St-Jacques (Vung Tau), 18 Mar 1875, Godefroy s.n. (lectotype K!, designated by Vidal and Hul Thol, 1976).

Caesalpinia thorelii Gagnep., Notul. Syst. (Paris). 2: 207. 1912.

Types. VIETNAM, 1^er pont de l’avalanche près Saïgon, 14 Jan 1865, Lefèvre, Thorel et Godefroy no. 145 (syntype P02940578!); Cochinchine, Bien-hoa, Nov 1866, Thorel 848 (syntype P02940348!); ad Bienhoa, Pierre 130 (syntype P02940353); Cochinchine, Baria, Baudoin and Talmy 104 (syntype);

Caesalpinia millettii Hook. & Arn., Bot. Beechey Voy. 182 (1841[1833]).

CHINA, Millett s.n. (K!).

Pterolobium subvestitum Hance, J. Bot. 22(12): 365. 1884.

Cantuffa subvestita (Hance) Kuntze, Rev. Gen. Pl. 1: 168. 1891.

Type. CHINA, Kwangtung, Lo Fau Sahn, Faber in herb. Hance 22291 (BM).

Caesalpinia oppositifolia Hattink, Reinwardtia 9(1): 43. 1974.

MALESIA, Sabah [North Borneo], Ranau Distr. Hot Spring track, 15 Feb 1961, J. Singh 24026 (holotype SAN; isotypes K!, L).

Caesalpinia parviflora Prain ex King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 66: 230. 1897.

MALAY PENINSULA, Perak, Relau Tugor, May 1888, Wray 1909 (lectotype CAL, designated by Hattink 1974; isolectotypes K!, SING).

Caesalpinia parviflora var. stipularis Prain, in J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 66: 230. 1897.

Types. MALAY PENINSULA, Perak, Larut, Wray 3983, 3991, 4261 (syntypes).

Caesalpinia stipularis Ridl., in Fl. Malay Penin. 1: 651 (1922), nom. illeg., non Caesalpinia stipularis (Vogel) Benth. (1870) (= Pomaria stipularis (Vogel) B.B. Simpson & G. P. Lewis).

Caesalpinia parviflora var. typica (Prain ex King) Prain, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 60: 230. 1897, nom. illeg.

Caesalpinia borneensis Merr., Univ. Calif. Publ. Bot. 15: 104. 1929.

Type. BORNEO, Tawao, Elphinstone Prov., Oct 1922– Mar 1923, Elmer 21449 (holotype MO; isotypes A, BM, BO, K!, L, NY, P, SING, U, UC).

Caesalpinia macra Craib, Bull. Misc. Inform. Kew 2: 386. 1927.

Type. THAILAND, Saraburi, Muak Lek, 10 Nov 1924, Marcan 1866 (syntype K), Pak Chong, 30 Dec 1923, Marcan 1532 (syntype K).

Caesalpinia minutiflora Elmer, Leafl. Philipp. Bot. 5: 1803. 1913.

Type. PHILIPPINES, Palawan, Puerto Princesa, Mt. Pulgar, Apr 1911, Elmer 12969 (BM, K!, L, P, PNH, U).

Caesalpinia sappan L., Sp. Pl. 1: 381. 1753.

SRI LANKA (CEYLON), Hb. Hermann, vol. 4, fol. 31 (holotype BM).

Caesalpinia angustifolia Salisb., Prod.: 326. 1796, nom. illeg.

Pterolobium lacerans R. Br. ex Wight & Arn., nom. illeg. (Cantuffa exosa J.F. Gmel. = Pterolobium exosum (J.F. Gmel.) E.G. Baker; this now considered a synonym of Pterolobium stellatum (Forssk.) Brenan).

Lianas or scrambling / trailing shrubs, armed with prickles on shoots, as well as in pairs at the base of leaves. Stipules small, inconspicuous, subulate or triangular-subulate, caducous. Leaves alternate, bipinnate, ending in a pair of pinnae, 6–30 cm long; petiole and rachis pubescent to sparsely pubescent or glabrous; pinnae opposite, in 5–20 pairs; leaflets opposite, in 6–25 pairs per pinna, linear-oblong to elliptic-oblong, apex rounded to emarginate, sometimes mucronate, eglandular or punctate-glandular, 6–15 × 1.5–10 mm. Inflorescences terminal or axillary racemes, often aggregated into panicles, pubescent to glabrous, 4–25 cm long; bracts small, caducous. Flowers bisexual, sub-actinomorphic to zygormophic; calyx comprising a short hypanthium and 5 sepals, glabrous to pubescent, the lower sepal cucullate, covering the other 4 sepals in bud; petals 5, free, yellow to white, equal to slightly differentiated, claws pubescent, the median petal sometimes inrolled; stamens 10, free, filaments pubescent (occasionally glabrous); ovary pubescent, stigma chambered. Fruit a red to brown samara, the basal seed-containing portion 12–20 × 8–15 mm, reticulate or smooth, glabrous to pubescent, the upper suture much prolonged and broadly winged, the wing 20–45 mm long and usually wider distally, 1 (–2)-seeded.

A genus of 10 species; one in southern tropical Africa, East Africa and Arabia, nine in SE Asia (one endemic to India, two in China, four in Indo-China [one endemic to Thailand, two extending to Malesia], three restricted to the Malay Peninsula and Archipelago [one endemic to the Philippines]).

Seasonally dry tropical upland evergreen forest, riverine and humid forest, woodland and wooded grassland.

From ptero- (Greek: wing) and lobion (Greek: pod, fruit), in reference to the fruit which is a samara.

Vidal and Hul Thol (1974) published a revision of Pterolobium, with a key to species. We provide below a list of species currently accepted in the genus, taking into account the treatment of P. sinense as a synonym of P. macropterum (Chen et al. 2010b).

Roti-Michelozzi (1957); Brenan (1967: 40–42); Vidal and Hul Thol (1974, 1976); Hul Thol and Hideux (1977); Hou et al. (1996: 654–700); Chen et al. (2010b).

Figure 23. 

Pterolobium stellatum (Forssk.) Brenan. A part of flowering branch B flower C longitudinal section of flower D petal E stamen F infructescence with mature fruits G samara with part cut away to reveal seed. A–E from Richards 11275 F from Eggeling 3400 G from Sandwith 25. Drawn by L. M. Ripley, originally published in Flora of Tropical East Africa, Leguminosae subfamily Caesalpinioideae, page 41, fig. 7 (1967).

Figure 24. 

Pterolobium stellatum (Forssk.) Brenan. A inflorescences (P. van Wyk, Africa, unvouchered) B fruits (J. Anton-Smith, Africa, unvouchered) C close up of fruits (B. T. Wursten, Flora of Zimbabwe (http://www.zimbabweflora.co.zw/speciesdata/image-display.php?species_id=127190&image_id=1), Zimbabwe, unvouchered). Mezoneuron hildebrandtii Vatke D inflorescences (D. Du Puy, Majunga, Madagascar, Du Puy M286 (P)) E fruits (D. Du Puy, Antsiranana, Madagascar, Du Puy M273 (P)). Mezoneuron kauaiense (H. Mann) Hillebr. F flower and buds I fruit (D. Eickhoff, Wikicommons (https://commons.wikimedia.org/wiki/Category:Mezonevron_kavaiense) cultivated, Hawaii, U.S.A., unvouchered). Caesalpinia crista L. emend. Dandy & Exell (?Ticanto) G flowers H young fruits (P. Grard: Institut Français de Pondichéry, Andhra Pradesh, India, unvouchered).

Mezoneuron glabrum Desf. ≡ Mezoneuron pubescens Desf.

Scrambling shrubs or lianas, occasionally medium -sized trees (M. kauaiense) to 12 m, usually armed with recurved prickles on stem and leaves, rarely unarmed. Stipules very small, often caducous. Leaves alternate or occasionally opposite, bipinnate, ending in a pair of pinnae; pinnae opposite to sub-opposite, in (1–)2–18 pairs; leaflets opposite to alternate, in 1–15 pairs per pinna, elliptic, oblong, suborbicular to occasionally subrhombic, the base oblique, the apex obtuse to acute. Inflorescences terminal or axillary racemes (often aggregated into panicles); bracteoles small. Flowers bisexual, zygomorphic; calyx comprising a hypanthium and 5 imbricate sepals, the lower sepal cucullate, and overlapping the other 4 in bud; petals 5, free, usually yellow with red markings on the median petal, or occasionally red, pink or cream, the median petal somewhat modified (either with a fleshy ligule or a patch of hairs on the inner surface between the blade and claw, or the petal bilobed); stamens 10, free, filaments alternately longer and shorter, usually all 10 pubescent or villous on lower half, or one or all glabrous; ovary glabrous to hairy, 1-many ovuled, stigma cupular, funnel-shaped, terminal or laterally placed, glabrous, or the rim fimbriate with papillate hairs, not peltate. Fruit laterally compressed, indehiscent, chartaceous, coriaceous or woody, venose, longitudinally and often broadly winged along the upper suture, the wing 1–18 mm wide. Seeds 1–13 per pod, ± transversely arranged in seed chamber, compressed, endosperm lacking.

A genus of 24 extant species, mainly in Asia, extending to Australia, Polynesia, Madagascar and Africa; two species on mainland Africa (one widespread in West Africa, the other in both West, East and Southeast Africa); one endemic to Madagascar; five endemic to New Caledonia; one endemic in Hawaii; one in Vietnam; four endemic to Australia (Queensland and New South Wales); one endemic in the Philippines; one in Australia and Papua New Guinea; nine species more widespread across Asia.

Tropical and subtropical riverine forest, lowland rain forest, swamp forest, seasonally dry forest, thicket, vine forest and wooded grassland, especially along forest and river margins.

From meso- (Greek: middle) or meizon (Greek: greater) and neuron (Greek: nerve), the upper suture of the fruit is bordered by a usually broad longitudinal wing so that the suture appears as a prominent sub-central nerve or vein.

The genus has recently been revised by Clark (2016), who provides full synonymy, a key to species, and a list of fossil taxa associated with this genus.

Brenan (1967: 38–40); Hattink (1974); Vidal and Hul Thol (1976); Verdcourt (1979: 18–20); Lock (1989: 25); Herendeen and Zarucchi (1990); Pedley (1997); George (1998: 59–67); Wagner et al. (1999); Du Puy and Rabevohitra (2002: 48–49); Brummitt et al. (2007); Clark and Gagnon (2015); Clark (2016).

Figure 25. 

Mezoneuron scortechinii F. Muell. A flowering branch B bract C calyx opened out D median petal E upper lateral petal F lower lateral petal G stamen H gynoecium I stigma J fruit K seed L detail of prickle from leaf. A–I, L from Hoogland 11665 J from Thurtill & Coveny 3880 K from White s.n. 6/1926. Drawn by Eleanor Catherine.